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Predicting tree frost tolerance is critical to select adapted species according to both the current and predicted future climate. The relative change in water to carbohydrate ratio is a relevant trait to predict frost acclimation in branches from many tree species. The objective of this study is to demonstrate the interspecific genericity of this approach across nine tree species. In the studied angiosperm species, frost hardiness dynamics were best correlated to a decrease in water content at the early stage of acclimation (summer and early autumn). Subsequently, frost hardiness dynamics were more tightly correlated to soluble carbohydrate contents until spring growth resumption. Based on different model formalisms, we predicted frost hardiness at different clade levels (angiosperms, family, genus and species) with high to moderate accuracy (1.5–6.0 °C root mean squared error (RMSE)) and robustness (2.8–6.1 °C prediction RMSE). The TOT model, taking all soluble carbohydrate and polyols into account, was more effective and adapted for large scale studies aiming to explore frost hardiness across a wide range of species. The ISC model taking the individual contribution of each soluble carbohydrate molecule into account was more efficient at finer scale such as family or species. The ISC model performance also suggests that the role of solutes cannot be reduced to a ‘bulk’ osmotic effect as could be computed if all of them were located in a single, common, compartment. This study provides sets of parameters to predict frost hardiness in a wide range of species, and clues for targeting specific carbohydrate molecules to improve frost hardiness.

The interaction between carbon and flows within the plant is at the center of most growth and developmental processes. Understanding how these fluxes influence each other, and how they respond to heterogeneous environmental conditions, is important to answer diverse questions in forest, agriculture and environmental sciences. However, due to the high complexity of the plant-environment system, specific tools are needed to perform such quantitative analyses.Here we present CPlantBox, full plant modelling framework based on the root system model CRootBox. CPlantbox is capable of simulating the growth and development of a variety of plant architectures (root and shoot). In addition, the flexibility of CPlantBox enables its coupling with external modeling tools. Here, we connected it to an existing mechanistic model of water and carbon flows in the plant, PiafMunch.The usefulness of the CPlantBox modelling framework is exemplified in four case studies. Firstly, we illustrate the range of plant structures that can be simulated using CPlantBox. In the second example, we simulated diurnal carbon and water flows, which corroborates published experimental data. In the third case study, we simulated impacts of heterogeneous environment on carbon and water flows. Finally, we showed that our modelling framework can be used to fit phloem pressure and flow speed to (published) experimental data.The CPlantBox modelling framework is open-source, highly accessible and flexible. Its aim is to provide a quantitative framework for the understanding of plant-environment interaction.

International audience; An important group of fungal diseases, such as apple scab on Malus x domestica leaves, perform infection only by remaining wetted by liquid water for a sufficient time, making leaf wetness duration (LWD) a crucial variable in epidemiological risk assessment. LWD is affected by the leaf wettability, i.e. the geometry of a liquid on a leaf as a consequence of the interaction between molecules of water and structural and physicochemical properties of the leaf surface. The variables describing wettability are the contact angles, which result from a genotype's physical and chemical properties. Accordingly, we assumed that wettability could be a direct expression of the genetic resistance of apple genotypes, thus influencing the physicochemical properties of the surface. In order to verify that hypothesis we measured static and dynamic contact angles on seven different Malus x domestica genotypes with various apple scab susceptibilities. The measured contact angle hysteresis was very high (from 58.8° to 82.6° among genotypes), leading to a high variability in possible values of static contact angles of water drops on leaf surface. The results obtained by mimicking both dew like and rain like water depositions during experiments, emphasize the importance of the water deposition process on the static contact angle of a water droplet on a leaf. A positive correlation between dew like droplets and the apple cultivar susceptibility to apple scab was observed and is discussed. This study is valuable for new LWD modeling approaches taking into account the leaf wetting properties.

In agroforestry systems, shade trees strongly affect the physiology of the undergrown crop. However, a major paradigm is that the reduction in absorbed photosynthetically active radiation is, to a certain extent, compensated by an increase in light-use efficiency, thereby reducing the difference in net primary productivity between shaded and non-shaded plants. Due to the large spatial heterogeneity in agroforestry systems and the lack of appropriate tools, the combined effects of such variables have seldom been analysed, even though they may help understand physiological processes underlying yield dynamics. In this study, we monitored net primary productivity, during two years, on scales ranging from individual coffee plants to the entire plot. Absorbed radiation was mapped with a 3D model (MAESPA). Light-use efficiency and net assimilation rate were derived for each coffee plant individually. We found that although irradiance was reduced by 60% below crowns of shade trees, coffee light-use efficiency increased by 50%, leaving net primary productivity fairly stable across all shade levels. Variability of aboveground net primary productivity of coffee plants was caused primarily by the age of the plants and by intraspecific competition among them (drivers usually overlooked in the agroforestry literature) rather than by the presence of shade trees.

Modeling is a fundamental part of quantitative science used to bring together several quantitative components, often developed though detailed reductionist approach on component parts, e.g., sucrose transport through a membrane osmotic relation. It is now generally accepted that phloem transport is the result of bulk solution flow generated by the difference in osmotic pressure between source and sink tissues. However, there is still little agreement on how different sink tissues compete for available carbohydrate. Furthermore, the impact of phloem pathway leakage (unloading) and reloading on source-to-sink carbon transport remains unclear. Moreover, it is debated to what degree the interactions between phloem and xylem flows influence carbohydrate source-sink relations. These aspects are extremely difficult to research by a reductionist approach, with modeling being an important tool to examine the consequences of proposed mechanisms, which can then be tested on whole plants.Phloem/xylem modeling has been at the limits of quantitative modeling, especially when dynamic models are needed to explain tracer studies. Advances in computing now enable more realistic modeling, which are utilized by the PiafMunch approach described here. This model enables a high level of mechanistic detail to be incorporated and the observable effect of it to be tested. In the most recent version of the software with the introduction of tracer dynamics, it can now predict the effects of specific phloem mechanisms upon the shape of evolving tracer profiles.

Modeling is a fundamental part of quantitative science used to bring together several quantitative components, often developed though detailed reductionist approach on component parts, e.g., sucrose transport through a membrane osmotic relation. It is now generally accepted that phloem transport is the result of bulk solution flow generated by the difference in osmotic pressure between source and sink tissues. However, there is still little agreement on how different sink tissues compete for available carbohydrate. Furthermore, the impact of phloem pathway leakage (unloading) and reloading on source-to-sink carbon transport remains unclear. Moreover, it is debated to what degree the interactions between phloem and xylem flows influence carbohydrate source-sink relations. These aspects are extremely difficult to research by a reductionist approach, with modeling being an important tool to examine the consequences of proposed mechanisms, which can then be tested on whole plants.Phloem/xylem modeling has been at the limits of quantitative modeling, especially when dynamic models are needed to explain tracer studies. Advances in computing now enable more realistic modeling, which are utilized by the PiafMunch approach described here. This model enables a high level of mechanistic detail to be incorporated and the observable effect of it to be tested. In the most recent version of the software with the introduction of tracer dynamics, it can now predict the effects of specific phloem mechanisms upon the shape of evolving tracer profiles.

International audience

Plant growth and development are limited by the available resources. The carbon and water flows are both important content and indicators of resource translocations. Modelling is helpful to increase the spatial and temporal resolutions of carbon and water reallocations in plants. However, the mechanism of carbon and water translocations has not been coupled into a whole Functional-Structural Plant Model (FSPM) yet. Here we developed a FSPM called CPlantBox which could (1) simulate the growth and development of the full plant structure (both root and shoot); (2) connect to a mechanistic model of water and carbon flow (PiafMunch). Our results demonstrate how carbon and water are flowing inside a plant which has three sources (leaf) and two sinks (root). We anticipate the model can be used as a tool to explore the variabilities and possibilities of plant behavior. Furthermore, several tool sets will be developed to visualize the morphological and physiological attributes of plants, which are helpful to deepen our understanding of plants and produce more with less.

It is now accepted that the transport phloem, linking major sources and sinks, is leaky, and this leakage can be considerable. Hence for phloem transport to function over the long distances observed, a large fraction of this unloaded photosynthate must be reloaded. A fraction of this unloaded solute is used to maintain tissues surrounding the phloem, as well as being stored. Also, pathway unloading/reloading acts as a short-term buffer to source and sink changes. In this work we present the first attempt to include both pathway unloading and reloading of carbohydrate in the modelling of pressure driven flow to determine if this has any significant effect upon source–sink dynamics. Our results indicated that the flow does not follow Poiseuille dynamics, and that pathway unloading alters the solute concentration and hydrostatic pressure profiles. Hence, measurement of either of these without considerable other detail tells us very little about the flow mechanisms. With adequate reloading along the pathway, the effects of pathway unloading can completely compensate for, making the entire system look like one with no pathway unloading.

UMR AGAP - équipe AFEF - Architecture et fonctionnement des espèces fruitières; International audience; The onset of the growing season of trees has been earlier by 2.3 days per decade during the last 40 years in temperate Europe because of global warming. The effect of temperature on plant phenology is, however, not linear because temperature has a dual effect on bud development. On one hand, low temperatures are necessary to break bud endodormancy, and, on the other hand, higher temperatures are necessary to promote bud cell growth afterward. Different process-based models have been developed in the last decades to predict the date of budbreak of woody species. They predict that global warming should delay or compromise endodormancy break at the species equatorward range limits leading to a delay or even impossibility to flower or set new leaves. These models are classically parameterized with flowering or budbreak dates only, with no information on the endodormancy break date because this information is very scarce. Here, we evaluated the efficiency of a set of phenological models to accurately predict the endodormancy break dates of three fruit trees. Our results show that models calibrated solely with budbreak dates usually do not accurately predict the endodormancy break date. Providing endodormancy break date for the model parameterization results in much more accurate prediction of this latter, with, however, a higher error than that on budbreak dates. Most importantly, we show that models not calibrated with endodormancy break dates can generate large discrepancies in forecasted budbreak dates when using climate scenarios as compared to models calibrated with endodormancy break dates. This discrepancy increases with mean annual temperature and is therefore the strongest after 2050 in the southernmost regions. Our results claim for the urgent need of massive measurements of endodormancy break dates in forest and fruit trees to yield more robust projections of phenological changes in a near future.

Biomass equations are a helpful tool to estimate the tree and stand biomass production and standing stock. Such estimations are of great interest for science but also of great importance for global reports on the carbon cycle and the global climate system. Even though there are various collections and generic meta-analyses available with biomass equations for mature trees, reports on biomass equations for juvenile trees (seedlings and saplings) are mainly missing. Against the background of an increasing amount of reforestation and afforestation projects and forests in young successional stages, such equations are required. In this study we have collected data from various studies on the aboveground woody biomass of 19 common tree species growing in Europe. The aim of this paper was to calculate species-specific biomass equations for the aboveground woody biomass of single trees in dependence of root-collar-diameter (RCD), height (H) and the combination of the two (RCD2 H). Next to calculating species-specific biomass equations for the species available in the dataset, we also calculated generic biomass equations for all broadleaved species and all conifer species. The biomass equations should be a contribution to the pool of published biomass equations, whereas the novelty is here that the equations were exclusively derived for young trees. We thank the national research project “Ecosystem Services of Natural Forests at Forestry and Climate Policy (FKZ 3511 84 0200),” from the Federal Agency for Nature Conservation (BfN) of the Federal Ministry for the Environment Nature Conservation and Nuclear Safety (BMU) for funding this project.

Adequate radial water transport between elastic bark tissue and xylem is crucial in trees, because it smoothens abrupt changes in xylem water potential, greatly reducing the likelihood of suffering dangerous levels of embolism. The radial hydraulic conductance involved is generally thought to be constant. Evidence collected about variable root and leaf hydraulic conductance led us to speculate that radial hydraulic conductance in stem/branches might also be variable and possibly modulated by putative aquaporins. We therefore correlated diameter changes in walnut (Juglans regia L.) with changes in water potential, altered by perfusion of twig samples with d-mannitol solutions having different osmotic potentials. Temperature and cycloheximide (CHX; a protein synthesis inhibitor) treatments were performed. The temperature response and diameter change inhibition found in CHX-treated twigs underpinned our hypothesis that radial hydraulic conductance is variable and likely mediated by a putative aquaporin abundance and/or activity. Our data demonstrate that radial water transport in stem/branches can take two routes in parallel: an apoplastic and a cell-to-cell route. The contribution of either route depends on the hydraulic demand and is closely linked to a boost of putative aquaporins, causing radial conductance to be variable. This variability should be considered when interpreting and modelling diameter changes.

As observed for most stresses, tree frost resistance can be split into two main processes: avoidance and tolerance. Avoidance of freezing is achieved by introducing species only in the climatic context in which the probability of freezing events is very low for the sensitive stages of buds or stems; i.e., when good synchronism exists between the annual cycle and the critical climatic periods. Buds become able to grow only after chilling requirements have been satisfied (endodormancy released) during winter; they subsequently break after heat requirements have been completed (end of ecodormancy) in early spring. Actually, this period is often subject to more or less severe freezing events. Trees are also able to adjust their freezing tolerance by increasing their capacity of extracellular freezing and decreasing the possibility of intracellular freezing through the process of frost acclimation. Both freezing resistance processes (avoidance and tolerance) are environmentally driven (by photoperiod and temperature), but there are also genotypic effects among species or cultivars. Here, we evaluated the degree to which differences in dormancy release and frost acclimation were related to environmental and genetic influences by comparing trees growing in common garden conditions. This investigation was carried out for two winters in lowland and mountain locations on different walnut genotypes differing significantly for budburst dates. Chilling requirement for endodormancy release and heat requirement during ecodormancy were evaluated in all situations. In addition, frost acclimation was assessed by the electrolyte leakage method on stems from the same trees before leaf fall through budburst. No significant differences were observed in chilling requirements among genotypes. Moreover, frost acclimation dynamics were similar between genotypes or locations when expressed depending on chilling units accumulated since 15 September as a time basis instead of Julian day. The only exception was for maximal frost hardiness observed during winter with the timber-oriented being significantly more resistant than fruit-oriented genotypes. Heat requirement was significantly different among genotypes. Thus, growth was significantly faster in fruit-oriented than in wood-oriented genotypes. Furthermore, among wood-oriented genotypes, differences in growth rate were observed only at cold temperatures. Frost acclimation changes differed significantly between fruit- and wood- walnuts from January through budburst. In conclusion, from September through January, the acclimation dynamic was driven mainly by environmental factors whereas from January through budburst a significant genotype effect was identified in both frost tolerance and avoidance processes.

When the current level of carbohydrates produced by photosynthesis is not enough to meet the C demand for maintenance, growth or metabolism, trees use stored carbohydrates. In rubber trees (Hevea brasiliensis Muell. Arg.), however, a previous study (Silpi U., A. Lacointe, P. Kasemsap, S. Thanisawanyangkura, P. Chantuma, E. Gohet, N. Musigamart, A. Clement, T. Ameglio and P. Thaler. 2007. Carbohydrate reserves as a competing sink: evidence from tapping the rubber tree. Tree Physiol. 27: 881-889) showed that the additional sink created by latex tapping results not in a decrease, but in an increase in the non-structural carbohydrate (NSC) storage in trunk wood. In this study, the response of NSC storage to latex tapping was further investigated to better understand the trade-off between latex regeneration, biomass and storage. Three tapping systems were compared to the untapped Control for 2 years. Soluble sugars and starch were analyzed in bark and wood on both sides of the trunk, from 50 to 200 cm from the ground. The results confirmed over the 2 years that tapped trees stored more NSC, mainly starch, than untapped Control. Moreover, a double cut alternative tapping system, which produced a higher latex yield than conventional systems, led to even higher NSC concentrations. In all tapped trees, the increase in storage occurred together with a reduction in trunk radial growth. This was interpreted as a shift in carbon allocation toward the creation of reserves, at the expense of growth, to cover the increased risk induced by tapping (repeated wounding and loss of C in latex). Starch was lower in bark than in wood, whereas it was the contrary for soluble sugars. The resulting NSC was twice as low and less variable in bark than in wood. Although latex regeneration occurs in the bark, changes related to latex tapping were more marked in wood than in bark. From seasonal dynamics and differences between the two sides of the trunk in response to tapping, we concluded that starch in wood behaved as the long-term reserve compartment at the whole trunk level, whereas starch in bark was a local buffer. Soluble sugars behaved like an intermediate, ready-to-use compartment in both wood and bark. Finally, the dynamics of carbohydrate reserves appears a relevant parameter to assess the long-term performance of latex tapping systems.

Water yam (Dioscorea alata L.) is one of the most Popular tuber crops in the tropics. Although it is well known that photoperiod greatly affects yam development, little information is available on the effect of temperature on different developmental phases, and there has been no attempt to model the effect of both environmental factors. The aim of this study was to assess the combined effect of photoperiod and temperature on the development of two early maturing varieties of yam with similar growth pattern. For this We used a model proposed for potatoes grown under tropical conditions. Experimental information was obtained from 15 field experiments carried out in Guadeloupe (French West Indies) covering a wide range of planting dates. Two yam phases were analyzed: from emergence (EM) to tuber initiation (TI), and from TI to harvest (HA). The EM-HA period varied from 3 to 6 months, with the longer cycles corresponding to early planting dates (e.g. April). On average, EM-TI represented one-third of the EM-HA period, and was mainly affected by photoperiod and to a lesser extent by temperature. Both factors also affected the duration of TI-HA but their effects were less noticeable. The observed mean temperature during TI-HA was near the estimated optimum and its effect was less than that of photoperiod. The variation of the phase duration was higher for EM-TI (CV 45%) than for TI-HA (8%), which was satisfactorily explained by the model, For EM-TI there was a positive interaction between both environmental factors which, together with the greater influence of climatic conditions, resulted in much variability in its duration. For TI-HA, a small influence of environmental factors, coupled with a compensatory effect between environmental factors, resulted in a relatively smaller variation in the duration of this phase. The model estimated satisfactorily the dates and the duration of each phase. The root mean square error (RMSE) was 7.4 d which corresponded to 13% and 7% of the observed mean duration of EM-TI and TI-HA, respectively. Our results showed that small changes in photoperiod and temperature, which are very usual in the tropics, have a big effect on the tested yam varieties. We also showed that the model applied in this study can be a useful tool to predict yam development for management purposes as well as for the modeling of yam growth. (C) 2009 Elsevier B.V. All rights reserved.

[EN] Non-structural carbohydrates (NSC) in plant tissue are frequently quantified to make inferences about plant responses to environmental conditions. Laboratories publishing estimates of NSC of woody plants use many different methods to evaluate NSC. We asked whether NSC estimates in the recent literature could be quantitatively compared among studies. We also asked whether any differences among laboratories were related to the extraction and quantification methods used to determine starch and sugar concentrations. These questions were addressed by sending sub-samples collected from five woody plant tissues, which varied in NSC content and chemical composition, to 29 laboratories. Each laboratory analyzed the samples with their laboratory-specific protocols, based on recent publications, to determine concentrations of soluble sugars, starch and their sum, total NSC. Laboratory estimates differed substantially for all samples. For example, estimates for Eucalyptus globulus leaves (EGL) varied from 23 to 116 (mean = 56) mg g(-1) for soluble sugars, 6-533 (mean = 94) mg g-1 for starch and 53-649 (mean = 153) mg g-1 for total NSC. Mixed model analysis of variance showed that much of the variability among laboratories was unrelated to the categories we used for extraction and quantification methods (method category R-2 = 0.05-0.12 for soluble sugars, 0.10-0.33 for starch and 0.01-0.09 for total NSC). For EGL, the difference between the highest and lowest least squares means for categories in the mixed model analysis was 33 mg g-1 for total NSC, compared with the range of laboratory estimates of 596 mg g-1. Laboratories were reasonably consistent in their ranks of estimates among tissues for starch (r = 0.41-0.91), but less so for total NSC (r = 0.45-0.84) and soluble sugars (r = 0.11-0.83). Our results show that NSC estimates for woody plant tissues cannot be compared among laboratories. The relative changes in NSC between treatments measured within a laboratory may be comparable within and between laboratories, especially for starch. To obtain comparable NSC estimates, we suggest that users can either adopt the reference method given in this publication, or report estimates for a portion of samples using the reference method, and report estimates for a standard reference material. Researchers interested in NSC estimates should work to identify and adopt standard methods., M.G.R. was funded by McMaster fellowship (1158.C). S.P. was funded by Juan de la Cierva contract (MCI project) and project ARBALMONT/786-2012 (OPAN, MAAMA, Spain). F.P. was funded by Fondecyt 11121175. U.N. and M.T. were funded by the Estonian Ministry of Education and Science, grant IUT-8-3. N.G.M. and L.T.D. were funded by DOE-BER. H.D.A. was funded by LANL-LDRD. J.M.-V. was funded by the Spanish Government (CGL 2010-16376). S.H. was funded by the Montana Institute on Ecosystems' Graduate Enhancement Award from NSF EPSCoR Track-1 NSF-IIA-1443108. Valuable comments from Dr Mauricio Mencuccini (University of Edinburgh), Dan Binkley (Colorado State University) and two anonymous reviewers were also greatly appreciated.

Xylem vessels of Prunus persica Batsch (peach) and Juglans regia L. (walnut) are vulnerable to frost-induced embolism. In peach, xylem embolism increased progressively over the winter, reaching a maximum of 85% loss of hydraulic conductivity (PLC) in early March. Over winter, PLC in walnut approached 100%, but the degree of xylem embolism varied during the winter, reflecting the ability of walnut to generate positive xylem pressures in winter and spring. In contrast, positive xylem pressures were not observed in peach. Controlled freeze-thaw experiments showed that frost alone is insufficient to increase embolism in peach; evaporative conditions during thawing are also required. However, when both species were protected from frost, PLC was zero. At bud break, there was complete recovery from embolism in walnut, whereas PLC remained high in peach. Three mechanisms responsible for the restoration of branch hydraulic conductivity were identified in walnut: the development of stem pressure, the development of root pressure and the formation of a new ring of functional xylem, whereas only one mechanism was observed in peach (new functional ring). The climatic conditions necessary for the manifestation of these mechanisms were investigated.

To improve source–sink relationship based carbon-allocation models, the basic proportional model was extended to account for a well-known effect of individual source to sink distances: among different sinks of similar characteristics, the more distant from the source, the lower the allocation coefficient. This was achieved through multiplication of the sink strength value by a coefficient that is proportional to a decreasing, simple function of distance, f; the power form was chosen for both simplicity and theoretical reasons. The resulting model was parameterized and evaluated on the empirical allocation matrix of the ECOPHYS model, after grouping together several individual, small sinks of similar nature and close location to remove any phyllotaxy-related bias. Both goodness of fit and predictive value were significantly improved compared with the basic proportional model (f = constant). The f-extended model yielded even better results if segments of different nature or age on the source to sink pathway were assigned different weights in the expression of distance, whereas the default expression of f, with an exponent of –1 and no additive constant, was optimal with no further parameter required. Thus, only 7 parameters (3 for pathway segment weights and 4 for sink strength values) were sufficient to retrieve the original 68 independent experimental allocation coefficients with a reasonable degree of accuracy. Pathway segment weights likely reflect both intrinsic transport pathway properties and situation within the plant architecture; this is discussed in relation to the possibilities of generalization and practical use of the model.

The onset of the growing season of trees has been globally earlier by 2.3 days/decade during the last 50 yearsbecause of global warming and this trend is predicted to continue according to climate forecast. The effect oftemperature on plant phenology is however not linear because temperature has a dual effect on bud development.On one hand, low temperatures are necessary to break bud dormancy, and on the other hand higher temperaturesare necessary to promote bud cells growth afterwards. Increasing phenological changes in temperate woodyspecies have strong impacts on forest trees distribution and productivity, as well as crops cultivation areas.Accurate predictions of trees phenology are therefore a prerequisite to understand and foresee the impacts ofclimate change on forests and agrosystems.Different process-based models have been developed in the last two decades to predict the date of budburstor flowering of woody species. They are two main families: (1) one-phase models which consider only the ecodormancyphase and make the assumption that endodormancy is always broken before adequate climatic conditionsfor cell growth occur; and (2) two-phase models which consider both the endodormancy and ecodormancy phasesand predict a date of dormancy break which varies from year to year. So far, one-phase models have been able topredict accurately tree bud break and flowering under historical climate. However, because they do not considerwhat happens prior to ecodormancy, and especially the possible negative effect of winter temperature warmingon dormancy break, it seems unlikely that they can provide accurate predictions in future climate conditions. Itis indeed well known that a lack of low temperature results in abnormal pattern of bud break and developmentin temperate fruit trees. An accurate modelling of the dormancy break date has thus become a major issue inphenology modelling.Two-phases phenological models predict that global warming should delay or compromise dormancy breakat the species equatorward range limits leading to a delay or even impossibility to flower or set new leaves. Thesemodels are classically parameterized with flowering or budburst dates only, with no information on the dormancybreak date because this information is very scarce. We evaluated the efficiency of a set of process-based phenologicalmodels to accurately predict the dormancy break dates of four fruit trees. Our results show that modelscalibrated solely with flowering or budburst dates do not accurately predict the dormancy break date. Providingdormancy break date for the model parameterization results in much more accurate simulation of this latter, withhowever a higher error than that on flowering or bud break dates. But most importantly, we show also that modelsnot calibrated with dormancy break dates can generate significant differences in forecasted flowering or budbreak dates when using climate scenarios. Our results claim for the urgent need of massive measurements of dormancybreak dates in forest and fruit trees to yield more robust projections of phenological changes in a near future.

International audience; Twenty-seven individual tree growth models are reviewed. The models take into account the same main physiological processes involved in carbon metabolism (photosynthate production, respiration, reserve dynamics, allocation of assimilates and growth) and share common rationales that are discussed. It is shown that the spatial resolution and representation of tree architecture used mainly depend on model objectives. Beyond common rationales, the models reviewed exhibit very different treatments of each process involved in carbon metabolism. The treatments of all these processes are presented and discussed in terms of formulation simplicity, ability to account for response to environment, and explanatory or predictive capacities. Representation of photosynthetic carbon gain ranges from merely empirical relationships that provide annual photosynthate production, to mechanistic models of instantaneous leaf photosynthesis that explicitly account for the effects of the major environmental variables. Respiration is often described empirically as the sum of two functional components (maintenance and growth). Maintenance demand is described by using temperature-dependent coefficients, while growth efficiency is described by using temperature-independent conversion coefficients. Carbohydrate reserve pools are generally represented as black boxes and their dynamics is rarely addressed. Storage and reserve mobilisation are often treated as passive phenomena, and reserve pools are assumed to behave like buffers that absorb the residual, excessive carbohydrate on a daily or seasonal basis. Various approaches to modelling carbon allocation have been applied, such as the use of empirical partitioning coefficients, balanced growth considerations and optimality principles, resistance mass-flow models, or the source-sink approach. The outputs of carbon-based models of individual tree growth are reviewed, and their implications for forestry and ecology are discussed. Three critical issues for these models to date are identified: (i) the representation of carbon allocation and of the effects of architecture on tree growth is Achilles' heel of most of tree growth models; (ii) reserve dynamics is always poorly accounted for; (iii) the representation of below ground processes and tree nutrient economy is lacking in most of the models reviewed. Addressing these critical issues could greatly enhance the reliability and predictive capacity of individual tree growth models in the near future.; Les modèles de croissance d'individus arbres basés sur le fonctionnement carboné : une évaluation critique. Vingt-sept modèles simulant la croissance d'arbres à l'échelle individuelle sont évalués. Ces modèles prennent en compte les principaux processus impliqués dans le métabolisme carboné (assimilation photosynthétique, respiration, dynamique des réserves, allocation des assimilats et croissance). Les concepts communs à tous ces modèles sont discutés. Il est montré que l'échelle d'espace et la représentation de l'architecture utilisées dépendent principalement des objectifs du modèle. Au-delà de concepts communs, les modèles évalués utilisent des représentations très différentes pour chacun des processus impliqués dans le métabolisme carboné. Les différentes représentations de ces processus sont présentées et discutées en termes de simplicité de formulation, de capacité à prendre en compte la réponse aux variables environnementales, et de capacités prédictives. La représentation des gains de carbone va de relations purement empirique calculant la production annuelle de photosynthétats jusqu'à des modèles de photosynthèse foliaire à bases mécanistes prenant explicitement en compte les effets des principales variables environnementales. La respiration est souvent décrite de façon empirique comme la somme de deux composantes (maintenance et croissance). La demande de maintenance est calculée à partir de coefficients dépendant de la température, alors que l'efficience de croissance est calculée à partir de coefficients de conversion indépendant de la température. Les réserves carbonées sont généralement représentées comme des boîtes noires, et leur dynamique est rarement prise en compte. La mise en réserve et l'utilisation des réserves sont souvent traitées comme des processus passifs, les réserves servant souvent de compartiment tampon absorbant les assimilats produits en excès sur une base journalière ou saisonnière. De nombreuses approches ont été utilisées pour modéliser l'allocation de carbone, telles que l'utilisation de coefficients d'allocation empiriques, l'application des principes de l'équilibre fonctionnel et d'optimisation, l'utilisation de schémas flux-résistance, ou des approches sources-puits. Les sorties des modèles simulant le bilan carboné et la croissance de plantes ligneuses à l'échelle individuelle sont présentées, et leurs implications en foresterie et en écologie sont discutées. Trois points particulièrement critiques actuellement pour ces modèles sont identifiés : (i) la représentation de l'allocation du carbone et des effets de l'architecture sur la croissance de l'arbre est le talon d'Achille de la majorité de ces modèles ; (ii) la dynamique des réserves est toujours faiblement représentée ; (iii) la représentation du fonctionnement racinaire et de la gestion des nutriments dans l'arbre est absente dans presque tous les modèles évalués. Une meilleure prise en compte de ces points critiques devrait fortement améliorer la fiabilité et les capacités prédictives des modèles de croissance d'arbres à l'échelle individuelle dans le futur.

Summary In winter and spring, walnut trees exhibit variations of sugar content in the vascular sap. According to their location, the vessel-associated cells (VACs, also called contact cells) could control nutrient exchanges between the storage parenchyma and the xylem vessels. According to the literature, the recovery of sap (influx) occurs at the VAC/vessel interface via an H+/sugar symport that depends on the transmembrane pH gradient generated by the plasma membrane H+-ATPase. The aim of this study was to investigate the ATPase activity, using a perfusion technique that allows the use of several effectors: carbonyl cyanide m-chlorophenylhydrazone (CCCP) and fusicoccin (FC). During winter, the uncoupler CCCP revealed a low pH gradient between the xylem vessels and the vessel-associated cells. Under these conditions, FC, an activator of the H+-ATPase, had no effect on the pH of the perfusion solution, suggesting that the enzyme could be lightly active. In contrast, close to bud break, a high pH gradient was revealed by the use of CCCP, and an acidification of the perfusion solution was observed in the presence of FC. Moreover, cytochemical investigation showed high activity of two respiratory enzymes located in mitochondria: NAD-dependent isocitrate dehydrogenase and succinate dehydrogenase. The hypothesis is that in spring this high respiratory activity of VACs provides a consequent increase in available ATP that can be utilized by the plasma membrane H+-ATPase.

International audience; SIMWAL (SIMulated WALnut) is a structural-functional tree model developed for single young walnut tree (Juglans sp.). It simulates the 3D structure dynamics of the tree, and biomass partitioning among its different organs, for a period ranging from a few months to several years, according to climatic conditions (temperature, radiation and air CO$_2$ concentration) and pruning. The aerial part of the tree is represented by axes split into growth units, inter-nodes, buds and leaves. The root system is described very coarsely by three compartments (taproot, coarse root and fine root). Only carbon-related physiological processes, i.e., radiation interception, photosynthesis, respiration, photosynthate allocation, and reserve storage and mobilisation are taken into account. Water and mineral nutrients are assumed to be optimal. We describe the model, and present preliminary tests of its ability to simulate tree architecture dynamics and carbon balance compared with field observations. Data requirements, and limits and improvements of the model are discussed.; SIMWAL : un modèle d'arbre structure-fonction simulant la croissance d'un noyer en fonction du climat et de la taille. SIMWAL est un modèle d'arbre structure-fonction développé pour le jeune noyer (Juglans sp.). Il simule l'évolution de la structure 3D de l'arbre et la répartition de la biomasse entre ses différents organes, pour des périodes allant de quelques mois à quelques années, en fonction des conditions climatiques (température, radiation, concentration dans l'air du CO$_2$) et des opérations de taille. La partie aérienne de l'arbre est représentée par des axes, eux-mêmes divisés en unités de croissance, entrenœuds, bourgeons et feuilles. Le système racinaire est plus grossièrement décrit en trois compartiments : pivot, racines moyennes et racines fines. Seuls les processus physiologiques relatifs au carbone sont pris en compte : interception lumineuse, photosynthèse, respiration, répartition carbonée, et stockage et mobilisation des réserves. L'eau et les éléments minéraux sont considérés à l'optimum. Nous décrivons en détail le modèle et présentons quelques tests préliminaires visant à vérifier sa capacité à simuler l'évolution de l'architecture d'un arbre et son bilan de carbone en comparaison à des observations de terrain. Le manque de certaines connaissances, les limites et les améliorations possibles du modèle sont discutés.

Tree Physiol. 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Oxford univ press Oxford; In the literature, frost hardiness (FH) studies in trees have often been restricted to one organ (buds, leaves, needles or twigs). To extend our knowledge and gain a unified view, FH differences between organs and tissues or throughout the life of the tree have to be characterized in relation to physiological changes. In this study, different organs and tissues of young potted and mature orchard walnut trees (Juglans regia L.) were compared for seasonal changes in FH during different years. FH was assessed using the electrolyte leakage method. Physiological parameters were concomitantly monitored focusing on two significant traits: water content (WC) and carbohydrate content (glucose + fructose + sucrose, GFS). No seasonal variation in FH was observed in the root system, but acclimation and deacclimation were observed aboveground. Among organs and tissues, cold sensitivity levels were different in deep winter, with buds most sensitive and bark most resistant, but acclimation/deacclimation dynamics followed similar patterns. Physiological variation was also similar among organs: FH increased when WC decreased and/or soluble carbohydrates increased. Based on these results, relations between soluble carbohydrate content, WC and FH were calculated independently or in interaction. The key results were that: (i) the relationship between FH and physiological parameters (GFS and WC), which had previously been shown for branches only, could be generalized to all aboveground organs; (ii) lower WC increased the cryoprotective effect of GFS, showing a synergic effect of the two factors; (iii) the best fit was a non-linear function of WC and GFS, yielding a predictive model with an root mean square error of 5.07 degrees C on an independent dataset and 2.59 degrees C for the most sensitive stages; and (iv) the same parameters used for all organs yielded a unified model of FH depending on physiology, although the variability of GFS or WC was wide. The model should be of value for predicting FH in walnut independently of previous growing conditions (i.e., after sublethal stress accumulation).

A closed growth chamber was designed to study the acquisition of autotrophy by seedlinps of walnut (Juglans regia L. cv. Lara) in controlled conditions (22 o C, 12-h photoperiod) during the first two months of growth. The chamber consisted of two airlight compartments, in which continuous gas exchange was measured on the aerial and subterranean parts of several batches of tree seedlings. Long-term labelling with 13 CO 2 was used in the chamber to study the import, distribution, and respiratory losses of photoassimilates (autotrophic carbon) in relation to the partitioning and use of reserves of the maternal seed (heterotrophic carbon)

Carbon (C) and nitrogen (N) autotrophy was studied in walnut seedlings (Juglans regia L.) grown in controlled conditions (22 o C, 12 h, 90% RH) during the first two months of development. Seedlings were submitted to simultaneous 13 CO 2 (1.3 13 C atom %) and 13 NO 3 (2.0 15 N atom %) long-term labelling, to quantify the assimilation and partitioning of recent C and N in growing organs compared to the mobilization and use of C and N derived from the seed reserves. The first organ to develop, at day 8, was the taproot, which built its biomass essentially during the first month with the help of C and N reserves of the seed

Adequate radial water transport between elastic bark tissue and xylem is crucial in trees, because it smoothens abrupt changes in xylem water potential, greatly reducing the likelihood of suffering dangerous levels of embolism. The radial hydraulic conductance involved is generally thought to be constant. Evidence collected about variable root and leaf hydraulic conductance led us to speculate that radial hydraulic conductance in stem/branches might also be variable and possibly modulated by putative aquaporins. We therefore correlated diameter changes in walnut (Juglans regia L.) with changes in water potential, altered by perfusion of twig samples with D-mannitol solutions having different osmotic potentials. Temperature and cycloheximide (CHX; a protein synthesis inhibitor) treatments were performed. The temperature response and diameter change inhibition found in CHX-treated twigs underpinned our hypothesis that radial hydraulic conductance is variable and likely mediated by a putative aquaporin abundance and/or activity. Our data demonstrate that radial water transport in stem/branches can take two routes in parallel: an apoplastic and a cell-to-cell route. The contribution of either route depends on the hydraulic demand and is closely linked to a boost of putative aquaporins, causing radial conductance to be variable. This variability should be considered when interpreting and modelling diameter changes.

Leaf Wetness Duration (LWD) is the main parameter involved in the apple scab development in apple orchards. LWD values can be inferred from weather conditions by solving a non-linear energy balance equation. Such energy balance equation is usually linearized leading to the Penman-Monteith equations. However this approximation leads to significant errors in the evaporative term and by consequence in the LWD computation. In this work, consequences of the use of the Penman-Monteith equations instead of the exact solution were firstly investigated on single droplets and compared to experimental values obtained in controlled and constant conditions. For these studies, the LWD is overestimated by 12% when Penman-Monteith equations were used. The ability of LWD estimate from the linearized energy balance equation to forecast risk infection of a leaf by V. inaequalis conidia was investigated. For some conditions, the error in LWD estimate by Penman-Monteith approached 6 h, leading to an incorrect risk assessment of apple scab development.

A mechanistic model of carbon partitioning, based on the Münch hypothesis of phloem transport and implemented with PIAF-Münch modelling platform (Lacointe and Minchin 2008), was tested for an architecture more complex than any tested previously. Using 11C to label photosynthate, responses in transport of photosynthate within a heavily pruned dwarf bean plant (Phaseolus vulgaris L.) to changes in source and sink activities were compared with model predictions. The observed treatment responses were successfully predicted. However, the observations could not be completely explained if the modelled stem contained only one phloem pathway: tracer from a labelled leaf was always detected in both shoot apex and root, whichever of the two leaves was labelled. This shows that bidirectional flow occurred within the stem, with solute moving simultaneously in both directions. Nevertheless, a model architecture with very little more complexity could incorporate such bidirectional flow. We concluded that the model could explain the observations, and that the PIAF-Münch model platform can be expected to describe partitioning in even more complex architectures.

Summary In 3-year-old walnut, a significant part of the starch reserves was converted into soluble sugars in the aerial parts and the fine roots in winter, but at the whole tree level most reserves were used in spring. The reserves stored in autumn were mobilized massively at the very beginning of bud break, and allocated for the most part to respiration. The summer reserves were mobilized to a lesser extent; they were for the most part incorporated in new tissues. The dynamics of reserve mobilization differed among organs. Older (2-year-old) reserves were mobilized to a very slight extent, although they remained available.

Ce systeme, utilise pour le marquage de noyers greffes de 3 ans (surface foliaire: 1,7 m 2 ), se compose d'une chambre d'assimilation et d'un dispositif d'injection de CO 2 a commande electronique permettant une regulation continue de la concentration en CO 2 . Ne comportant pas de dispositif de regulation thermique, il n'est utilise que pendant la matinee. Malgre une augmentation significative de la temperature au cours du marquage, la photosynthese est peu perturbee: le taux d'assimilation (pente des segments decroissants) reste regulier (.)

It has been hypothesized that the increase in temperature in this century could lead to an increase in frost damage to plant tissues. Several models have been proposed to describe the development of cold hardiness, but never taking into account extreme climatic and/or physiological events. Our results on walnut tree (Juglans regia L.) show that cold hardiness was best correlated with average daily temperatures minimal temperatures over the last 15 days before sampling ((T) over bar (min 15 days)), indicating that the freezing tolerance depended on the tree's climatic history. Moreover, this study also shows that the accumulation of sucrose and the water content (WC) decrease are an essential step towards cold hardiness. Thus, a simple linear model based on climatic ((T) over bar (min 15 days)) and physiological (soluble sugars, WC) explanatory variables was developed to predict the cold hardiness level in walnut stem at any time during the leafless period. Each of the three input variables can be assigned a specific role contributing to the simulated function, cold hardiness. The extent and robustness of this relation was assessed on extreme physiological events on walnut trees bearing three main branches. On each tree, one branch was defoliated to limit the local carbohydrate and transpiration, one was girdled to increase local carbohydrate and prevent carbohydrate export and the third one was kept untreated as control. As expected, these treatments impacted both local carbon reserves and WC in the stems born by each main branch in comparison with the control on the same tree. The impact of these treatments on stem's freezing tolerance, as evaluated by an electrolyte leakage method (LT(50)), confirmed the direct impact of soluble sugar and WC on cold hardiness over a wide range of carbohydrate and WC. This is discussed in relation to the branch autonomy theory for carbon but also for water during summer growth and winter periods. The present study demonstrates the importance of physiological parameters in the prediction of cold hardiness and proposes a way to model cold hardiness with extreme climatic and/or physiological events.

International audience; Background and Aims: In rain forests, sapling survival is highly dependent on the regulation of trunk slenderness (height/diameter ratio): shade-intolerant species have to grow in height as fast as possible to reach the canopy but also have to withstand mechanical loadings (wind and their own weight) to avoid buckling. Recent studies suggest that mechanosensing is essential to control tree dimensions and stability-related morphogenesis. Differences in species slenderness have been observed among rainforest trees; the present study thus investigates whether species with different slenderness and growth habits exhibit differences in mechanosensitivity. Methods: Recent studies have led to a model of mechanosensing (sum-of-strains model) that predicts a quantitative relationship between the applied sum of longitudinal strains and the plant's responses in the case of a single bending. Saplings of five different neotropical species (Eperua falcata, E. grandiflora, Tachigali melinonii, Symphonia globulifera and Bauhinia guianensis) were subjected to a regimen of controlled mechanical loading phases (bending) alternating with still phases over a period of 2 months. Mechanical loading was controlled in terms of strains and the five species were subjected to the same range of sum of strains. The application of the sum-of-strain model led to a dose-response curve for each species. Dose-response curves were then compared between tested species. Key Results: The model of mechanosensing (sum-of-strain model) applied in the case of multiple bending as long as the bending frequency was low. A comparison of dose-response curves for each species demonstrated differences in the stimulus threshold, suggesting two groups of responses among the species. Interestingly, the liana species B. guianensis exhibited a higher threshold than other Leguminosae species tested. Conclusions: This study provides a conceptual framework to study variability in plant mechanosensing and demonstrated interspecific variability in mechanosensing.

Evaluation of endodormancy release models in temperate fruit tree species has generally been indirect, as a component of the global evaluation of bud break-date models that include an endodormancy release module. Due to the lack of data, their direct evaluation based on the knowledge of the dynamics of bud dormancy states has been very succinct. Consequently, these models have not been optimized, except for the 'amount of chill units required to break endodormancy' taken as the genotype specific parameter. As part of different studies on bud state during the rest period, near 30 yearly dynamics of the dormancy state of 'Redhaven' peach vegetative buds under natural conditions have been recorded, based on the biological 'one node cuttings' test under standard conditions (25C). These data were used to optimize classic models (Weinberger-W; Utah-U, etc.) of endodormancy release in fruit tree species with temperature as the only input variable, in order to compare their goodness of fit and their predictive capacity. The fitted parameters were the 'amount of chill units required to break endodormancy', those of the temperature functions f(T) describing the rate of endodormancy development and, when relevant, the starting date of endodormancy release. The results and practical conclusions for predicting dormancy release date are presented. The optimization method used was the minimization of the mean square error (MSE) between the observed (biological test) and the computed dates of endodormancy release and was carried out using the nls module of R software. As main results: the optimization of the W model resulted in an increase of the standard threshold temperature; the endodormancy release effect of negative temperatures (up to freezing injury) in the optimized form of U model that was not taken into account in the standard form; the sharp transition from positive to negative f(T) values for the temperatures above 12C, in the optimized U model; and the fact that the model of Weinberger is a particular form of smoothed Utah model.

Bud break pattern is a key determinant of tree architecture. The mechanisms leading to the precedence of certain buds over the others are not yet fully explained, but the availability of soluble sugars may play a significant role, especially those in the xylem sap at the onset of the growing period. Here, we measured carbon availability in the different tissues (bud, xylem and bark). To assess the capacity of buds to use the xylem sap carbohydrates, the fluxes between xylem vessels and parenchyma cells, bark and buds of walnut (Juglans regia cv 'Franquette') were measured during the rest period until bud break. This uptake capacity varies according to the temperature, the sugar and the position on the branch of the fragment studied. Between December and March, in xylem tissues, the active component of sucrose uptake was predominant compared with diffusion (90% of the total uptake), whereas the active component accounted for more moderate amounts in buds (50% of the uptake). The active uptake of hexoses took place belatedly (April) in xylem. The flow rates between xylem vessels and buds increased 1 month before bud break and reached 2000 microg sucrose h(-)(1) g DW(-)(1). Fluxes seemed to depend on bud position on the branch. However, this study strongly suggests that they were mainly dependent on the sink strength of the buds and on the sink competition between bud, xylem parenchyma and bark.

† Background and Aims The major economic product of Hevea brasiliensis is a rubber-containing cytoplasm (latex), which flows out of laticifers (latex cells) when the bark is tapped. The latex yield is stimulated by ethylene. Sucrose, the unique precursor of rubber synthesis, must cross the plasma membrane through specific sucrose transporters before being metabolized in the laticifers. The relative importance of sucrose transporters in determining latex yield is unknown. Here, the effects of ethylene (by application of Ethrel w ) on sucrose transporter gene expression in the inner bark tissues and latex cells of H. brasiliensis are described. † Methods Experiments, including cloning sucrose transporters, real time RT-PCR and in situ hybridization, were carried out on virgin (untapped) trees, treated or untreated with the latex yield stimulant Ethrel. † Key Results Seven putative full-length cDNAs of sucrose transporters were cloned from a latex-specific cDNA library. These transporters belong to all SUT (sucrose transporter) groups and differ by their basal gene expression in latex and inner soft bark, with a predominance of HbSUT1A and HbSUT1B. Of these sucrose transporters, only HbSUT1A and HbSUT2A were distinctly increased by ethylene. Moreover, this increase was shown to be specific to laticifers and to ethylene application. † Conclusion The data and all previous information on sucrose transport show that HbSUT1A and HbSUT2A are related to the increase in sucrose import into laticifers, required for the stimulation of latex yield by ethylene in virgin trees.

absent

The function of the plant’s vasculature, incorporating both phloem and xylem, is of fundamental importance to the survival of all higher plants. Although the physiological mechanism involved in these two transport pathways has been known for some time, quantitative modelling of this has been slow to develop. 1-D continuous models have shown that the proposed mechanisms are quantitatively plausible (Thompson and Holbrook 2003) but more complex geometries (architectures) have remained out of reach because of mathematical difficulties. In this work, we extend the alternative modular approach by Daudet et al. (2002) using recently developed numerical tools which allow us to model complex architectures. After a full description of the extended model, we first show that it efficiently reproduces the results of the continuous approach when applied to the same simple configurations. The model is then applied to a more complex configuration with two sinks, confirming that sink priority is an emergent property of the Münch flow as earlier found with a minimalist model (Minchin et al. 1993). It is further shown how source leaf transpiration can change the relative carbon allocation rates among sinks.

International audience; The carbon economy of young walnut grafted trees in their second growing year, was investigated through carbohydrate analysis and 14C-labelling experiments. In October, 3 weeks before leaf-fall, starch and soluble sugars represented up to 25% of the total dry matter, with starch highly predominant among them. Sucrose was more abundant in the bark than in the wood, while the starch content was similar in both compartments. The reserves were located in the wood and in the innermost pith. Between October and January, the minor roots (ø < 15mm) underwent a significant depletion of their August-labelled reserves, whereas the tap-root was the main organ depleted between January and May (early growth resumption). Both labelling experiments, and root respiration monitoring, provided some information about the timing of translocation processes and the early metabolism of the translocation compounds.

International audience; Plasma membrane H+-ATPase (PM H+-ATPase) plays a key role in nutrient transport, stress responses and growth. To evaluate proton motive force differences between apical and basal parts of acrotonic 1-year-old shoots of walnut (Juglans regia L. cv 'Franquette') trees, spatial and seasonal changes in PM H+-ATPase were studied in mature xylem tissues. During both the dormancy and growth resumption periods, and in both the apical and basal parts of the stem, PM H+-ATPase activity showed positive correlations with the amount of immunodetectable protein. In spring, at the time of growth resumption, higher activities and immunoreactivities of PM H+-ATPase were found in the apical part of the stem than in the basal part of the stem. In spring, the decrease in xylem sugar concentration reflected the high sugar uptake rate. Our data suggest that PM H+-ATPase plays a major role in the uptake of carbohydrates from xylem vessels during growth resumption. These results are discussed in the context of the acrotonic tendency of walnut shoots.

Carbohydrate reserve storage in trees is usually considered a passive function, essentially buffering temporary discrepancies between carbon availability and demand in the annual cycle. Recently, however, the concept has emerged that storage might be a process that competes with other active sinks for assimilate. We tested the validity of this concept in Hevea brasiliensis Mull. Arg. (rubber) trees, a species in which carbon availability can be manipulated by tapping, which induces latex regeneration, a high carbon-cost activity. The annual dynamics of carbohydrate reserves were followed during three situations of decreasing carbon availability: control (no tapping), tapped and tapped with Ethephon stimulation. In untapped control trees, starch and sucrose were the main carbohydrate compounds. Total nonstructural carbohydrates (TNC), particularly starch, were depleted following bud break and re-foliation, resulting in an acropetal gradient of decreasing starch concentration in the stem wood. During the vegetative season, TNC concentration increased. At the end of the vegetative season, there were almost no differences in TNC concentration along the trunk. In tapped trees, the vertical gradient of starch concentration was locally disturbed by the presence of the tapping cut. However, the main effect of tapping was a dramatic increase in TNC concentration, particularly starch, throughout the trunk and in the root. The difference in TNC concentration between tapped and untapped trees was highest when latex production was highest (October); the difference was noticeable even in areas of the trees that are unlikely to be directly involved in latex regeneration, and it was enhanced by Ethephon stimulation, which is known to increase latex metabolism and flow duration. Thus, contrary to what could be expected if reserves serve as a passive buffer, a decrease in carbohydrate availability resulted in a net increase in carbohydrate reserves at the trunk scale. Such behavior supports the view that trees tend to adjust the amount of carbohydrate reserves stored to the level of metabolic demand, at the possible expense of growth.

This study describes the sucrose balance between supply and demand in the bark of the rubber tree, along with concurrent latex metabolic activity. Experiments were designed using three H. brasiliensis clones (PB 235, RRIM 600 and GT (1)) in the same polyclonal plot at the Chachoengsao Rubber Research Centre (CRRC-DOA) in Thailand. Treatments were carried out on previously untapped trees (growth potential control), trees tapped without stimulation (1/2 S d/3 6 d/7 9 m/12, physiological control), and trees tapped with ethephon stimulation (1/2 S d/3 6 d/7 9 m/12 ET 2.5% 5/y and 12/y). Tapping had a marked effect on latex physiology in the whole trunk. Sucrose concentration was significantly reduced. The Latex Diagnosis Mapping (LDM) method was used to describe the shape and size of the latex regeneration area and of the metabolically active bark area. For the three clones, rubber production correlated with the estimated latex regeneration area. It took around 100 cm2 of latex regeneration area to regenerate 1 g of rubber. As it assesses the impact of any tapping system on whole trunk latex physiology, the LDM method was used to develop new tapping systems, such as systems involving ethylene gas stimulation, micro-tapping cut systems, and multi-tapping cut systems.

The respective roles of carbon and water in radial growth were assessed using double girdling and water stress as experimental factors to induce variations of the carbon and water status of tissues of Juglans nigra x Juglans regia grown in France. The results show that structural radial growth (with its specific energy requirement) occurred mainly during the night-time, in relation with an improved water status, and consequently higher turgor pressure which allows tissue enlargement (cell division and/or expansion). Maximal respiration rate occurring at night, when transpiration was minimal, suggests no or very low involvement of dissolved CO2 originating from a distance, i.e. carried by the transpirational flux of xylem sap, in the total stem CO2 efflux rate. The observations point out the importance of current photoassimilate flux for active radial growth. Local stem reserves could only marginally contribute to radial growth, although sharp shortage in assimilate supply could induce, within a few days, significant mobilization of local parenchyma reserves which allowed growth resumption. However, the driving force of mobilization was apparently not organ-wide sugar deficiency, as mobilization was not associated with any significant drop of sugars at the organ level.

The variations of stem diameter as they can be accurately measured by Linear Variable Differential Transformer (LVDT) reflect the addition of four components: irreversible radial growth, reversible living-cell dehydration/rehydration, thermal expansion and contraction, and expansion of dead conducting elements due to the increase and relaxation of internal tensions. The correct interpretation of LVDT signals, with respect to the practical applications, should make an exact distinction between these four components. This paper describes a set of two experiments with potted hybrid walnut trees. Double girdling, water stress, and duration of the day versus night periods were used in the phytotron as experimental factors to induce variations of the carbon and water status of plant tissues. The latter were assessed, respectively, by water potential and transpiration, and by local stem respiration and carbohydrate content. The results are interpreted in terms of carbon or water limitation effects on stem diameter variations where radial growth and tissue elasticity could be distinguished. Moreover, they suggest no or very low involvement of CO2 originating from a distance, i.e. carried by the transpirational flux of xylem sap, in the total stem CO2 efflux rate.

Summary Most current models of assimilate carbohydrate partitioning are based on growth patterns observed under a range of experimental conditions, from which a set of empirical rules are derived to simulate partitioning. As a result, they are not good at extrapolating to other conditions; this requires a mechanistic approach, which only transport-resistance (TR) models currently provide. We examine an approach to incorporating recent progress in phloem physiology into the TR approach, which leads to a ‘minimalist’ Munch model of a branched system with competing sinks. In vivo whole-plant measurements have demonstrated that C-flow rates are dependent not only on the properties of the sink, but also on the properties of the whole transport system, and the detailed dynamics of this behaviour is mimicked by the proposed model. This model provides a sound theoretical framework for an unambiguous definition of sink and source strengths, with sink priority being an emergent property of the model. Further developments are proposed, some of which have already had limited application, to cope with the complexity of plants; the emphasis is on a modular approach, together with the importance of choosing the appropriate scale level for both structure and function. Whole-plant experiments with in vivo measurement of the phloem dynamics will be needed to help with this choice.

Endogenous carbohydrate contents and availability in structures near the meristematic zone (scales, cushion, part of stem) during winter with and without sufficient chilling were investigated. Insufficient chilling is known to provoke an abnormal pattern of budbreak and development in temperate zone fruit trees cultivated in warm climates. Mild temperatures applied during winter to strictly floral buds, as is the case in peach, are known to provoke the necrosis of flower primordia in the buds. One of the hypotheses to explain this is that carbohydrate fluxes to the buds are impeded and/or that internal reserves are completely used up. Carbohydrate availability and bud survival were compared between peach trees under natural temperate conditions and under complete deprivation of cold, by keeping trees in a greenhouse with temperature constantly above 15 °C from early October to the first sign of necrosis in flower buds. Concentrations and carbohydrate contents were compared during a little more than 2 months (beginning mid-December) before the first signs of necrosis in flower buds. Results showed that cold deprivation deeply disturbed carbohydrate dynamics, blocking the import of carbohydrates into vegetative and floral primordia, even though sucrose and sorbitol concentrations within these structures remained high. Conversely, soluble carbohydrates accumulated strongly in bud scales and in the cushion beneath the primordia. This suggests a strong diversion towards structures close to the primordia under cold deprivation. However, since sucrose and sorbitol concentrations in the floral primordia remained high, necrosis could be the consequence of the inability to use sucrose and sorbitol reserves rather than the result of a lack of available carbohydrates.

In south-central France, walnut exhibited freezing tolerance by acclimation in the fall and deacclimation in the spring. The cold hardiness level varied with the different cultivars and dates. Cold tolerance performances of 7 walnut fruit cultivars (Chandler, Franquette, Fernor, Fernette, Lara, Pedro, Serr) and of the hybrid Juglans regia × Juglans nigra NG38 were compared in the same location, at the INRA Center of Clermont-Ferrand. For these comparisons, an electrolyte leakage conductivity method (LT50) was used and adapted for walnut.In all cases NG38 (Juglans regia × Juglans nigra) appeared particularly cold resistant.