Dominique Gérant

Pulse-labelling trees to study carbon allocation dynamics: a review of methods, current knowledge and future prospects

Pulse-labelling of trees with stable or radioactive carbon (C) isotopes offers the unique opportunity to trace the fate of labelled CO(2) into the tree and its release to the soil and the atmosphere. Thus, pulse-labelling enables the quantification of C partitioning in forests and the assessment of the role of partitioning in tree growth, resource acquisition and C sequestration. However, this is associated with challenges as regards the choice of a tracer, the methods of tracing labelled C in tree and soil compartments and the quantitative analysis of C dynamics. Based on data from 47 studies, the rate of transfer differs between broadleaved and coniferous species and decreases as temperature and soil water content decrease. Labelled C is rapidly transferred belowground-within a few days or less-and this transfer is slowed down by drought. Half-lives of labelled C in phloem sap (transfer pool) and in mature leaves (source organs) are short, while those of sink organs (growing tissues, seasonal storage) are longer. (13)C measurements in respiratory efflux at high temporal resolution provide the best estimate of the mean residence times of C in respiratory substrate pools, and the best basis for compartmental modelling. Seasonal C dynamics and allocation patterns indicate that sink strength variations are important drivers for C fluxes. We propose a conceptual model for temperate and boreal trees, which considers the use of recently assimilated C versus stored C. We recommend best practices for designing and analysing pulse-labelling experiments, and identify several topics which we consider of prime importance for future research on C allocation in trees: (i) whole-tree C source-sink relations, (ii) C allocation to secondary metabolism, (iii) responses to environmental change, (iv) effects of seasonality versus phenology in and across biomes, and (v) carbon-nitrogen interactions. Substantial progress is expected from emerging technologies, but the largest challenge remains to carry out in situ whole-tree labelling experiments on mature trees to improve our understanding of the environmental and physiological controls on C allocation.

Seasonal changes of C and N non-structural compounds in the stem sapwood of adult sessile oak and beech trees

We assessed the pools of non-structural nitrogen compounds (NSNC) through a year, thereby addressing the question of whether mature sessile oak [Quercus petraea (Matt.) Liebl.] and beech (Fagus sylvatica L.), which differ in wood anatomy and growth patterns, exhibit contrasting seasonal dynamics of NSNC pools as previously shown for non-structural carbohydrate (NSC) pools. Seasonal fluctuations of NSNC (amino acids and soluble proteins) and NSC (starch and soluble sugars) pools were analyzed in the inner and the outer stem sapwood. In oak, NSC showed marked seasonal variation within the stem sapwood (accumulation during winter and decrease during bud burst and early wood growth), whereas in beech seasonal fluctuations in NSC were of minor amplitude. Even if the distribution and intensity of the NSNC pools differed between the two species, NSNC of the stem sapwood did not show seasonal variation. The most significant change in NSNC pools was the seasonal fluctuation of protein composition. In both species, two polypeptides of 13 kDa (PP13) and 26 kDa (PP26) accumulated during the coldest period in parallel with starch to sugar conversion and disappeared with the onset of spring growth. The absence of seasonal changes in total soluble protein concentration suggests that the polypeptides are involved in the internal nitrogen (N) cycling of the stem rather than in N storage and remobilization to the other growing organs of the tree.

Nitrogen sources for current-year shoot growth in 50-year-old sessile oak trees: an in situ 15N labeling approach

We used long-term in situ (15)N labeling of the soil to investigate the contribution of the two main nitrogen (N) sources (N uptake versus N reserves) to sun shoot growth from bud burst to full leaf expansion in 50-year-old sessile oaks. Recovery of (15)N by growing compartments (leaves, twigs and buds) and presence of (15)N in phloem sap were checked weekly. During the first 2 weeks following bud burst, remobilized N contributed ~90% of total N in growing leaves and twigs. Nitrogen uptake from the soil started concomitantly with N remobilization but contributed only slightly to bud burst. However, the fraction of total N due to N uptake increased markedly once bud burst had occurred, reaching 27% in fully expanded leaves and 18% in developed twigs. In phloem sap, the (15)N label appeared a few days after the beginning of labeling and increased until the end of bud burst, and then decreased at full leaf expansion in June. Of all the shoot compartments, leaves attracted most of the absorbed N, which accounted for 68% of new N in shoots, whereas twigs and new buds accounted for only 28 and 3%, respectively. New N allocated to leaves increased from unfolding to full expansion as total N concentration in the leaves decreased. Our results underline the crucial role played by stored N in rapid leaf growth and in the sustained growth of oak trees. Any factors that reduce N storage in autumn may therefore impair spring shoot growth.

In situ 13CO2 pulse labelling of field-grown eucalypt trees revealed the effects of potassium nutrition and throughfall exclusion on phloem transport of photosynthetic carbon

Potassium (K) is an important limiting factor of tree growth, but little is known of the effects of K supply on the long-distance transport of photosynthetic carbon (C) in the phloem and of the interaction between K fertilization and drought. We pulse-labelled 2-year-old Eucalyptus grandis L. trees grown in a field trial combining K fertilization (+K and -K) and throughfall exclusion (+W and -W), and we estimated the velocity of C transfer by comparing time lags between the uptake of (13)CO2 and its recovery in trunk CO2 efflux recorded at different heights. We also analysed the dynamics of the labelled photosynthates recovered in the foliage and in the phloem sap (inner bark extract). The mean residence time of labelled C in the foliage was short (21-31u2005h). The time series of (13)C in excess in the foliage was affected by the level of fertilization, whereas the effect of throughfall exclusion was not significant. The velocity of C transfer in the trunk (0.20-0.82u2005mu2005h(-1)) was twice as high in +K trees than in -K trees, with no significant effect of throughfall exclusion except for one +Ku2005-W tree labelled in the middle of the drought season that was exposed to a more pronounced water stress (midday leaf water potential of -2.2u2005MPa). Our results suggest that besides reductions in photosynthetic C supply and in C demand by sink organs, the lower velocity under K deficiency is due to a lower cross-sectional area of the sieve tubes, whereas an increase in phloem sap viscosity is more likely limiting phloem transport under drought. In all treatments, 10 times less (13)C was recovered in inner bark extracts at the bottom of the trunk when compared with the base of the crown, suggesting that a large part of the labelled assimilates has been exported out of the phloem and replaced by unlabelled C. This supports the leakage-retrieval mechanism that may play a role in maintaining the pressure gradient between source and sink organs required to sustain high velocity of phloem transport in tall trees.

Seasonal variations drive short‐term dynamics and partitioning of recently assimilated carbon in the foliage of adult beech and pine

13 CO2 pulse-labelling experiments were performed inxa0situ on adult beeches (Fagus sylvatica) and pines (Pinus pinaster) at different phenological stages to study seasonal and interspecific short-term dynamics and partitioning of recently assimilated carbon (C) in leaves. Polar fraction (PF, including soluble sugars, amino acids and organic acids) and starch were purified from foliage sampled during a 10-d chase period. C contents, isotopic compositions and 13 C dynamics parameters were determined in bulk foliage, PF and starch. Decrease in 13 C amount in bulk foliage followed a two-pool exponential model highlighting 13 C partitioning between mobile and stable pools, the relative proportion of the latter being maximal in beech leaves in May. Early in the growing season, new foliage acted as a strong C sink in both species, but although young leaves and needles were already photosynthesizing, the latter were still supplied with previous-year needle photosynthates 2 months after budburst. Mean 13 C residence times (MRT) were minimal in summer, indicating fast photosynthate export to supply perennial organ growth in both species. In late summer, MRT differed between senescing beech leaves and overwintering pine needles. Seasonal variations of 13 C partitioning and dynamics in field-grown tree foliage are closely linked to phenological differences between deciduous and evergreen trees.

Seasonality of nitrogen partitioning (non-structural vs structural) in the leaves and woody tissues of tropical eucalypts experiencing a marked dry season

Numerous studies have shown that internal nitrogen (N) translocation in temperate tree species is governed by photoperiod duration and temperature. For tropical tree species, the seasonality of rainfall is known to affect growth and foliage production, suggesting that efficient internal N recycling also occurs throughout the year. We tested this hypothesis by comparing the N budgets and N partitioning (non-structural vs structural N) in the different organs of 7-year-old Eucalyptus urophylla (S.T. Blake) × E. grandis (W. Hill ex Maiden) trees from a plantation in coastal Congo on poor sandy soil. The trees were sampled at the end of the dry season and late in the rainy season. Lower N concentrations and N investment in the non-structural fraction were observed in leaves during the dry season, which indicates resorption of non-structural N from senescing leaves. Stem wood, which contributes to about 60% of the total biomass of the trees, accumulated high amounts of non-structural N at the end of the dry season, most of which was remobilized during the following rainy season. These results support the hypothesis of efficient internal N recycling, which may be an important determinant for the growth potential of eucalypts on N-poor soils. Harvesting trees late in the rainy season when stem wood is depleted in non-structural N should be recommended to limit the export of nutrients off-site and to improve the sustainability of tropical eucalypt plantations.

The impact of prolonged drought on phloem anatomy and phloem transport in young beech trees

Phloem failure has recently been recognized as one of the mechanisms causing tree mortality under drought, though direct evidence is still lacking. We combined 13C pulse-labelling of 8-year-old beech trees (Fagus sylvatica L.) growing outdoors in a nursery with an anatomical study of the phloem tissue in their stems to examine how drought alters carbon transport and phloem transport capacity. For the six trees under drought, predawn leaf water potential ranged from -0.7 to -2.4 MPa, compared with an average of -0.2 MPa in five control trees with no water stress. We also observed a longer residence time of excess 13C in the foliage and the phloem sap in trees under drought compared with controls. Compared with controls, excess 13C in trunk respiration peaked later in trees under moderate drought conditions and showed no decline even after 4 days under more severe drought conditions. We estimated higher phloem sap viscosity in trees under drought. We also observed much smaller sieve-tube radii in all drought-stressed trees, which led to lower sieve-tube conductivity and lower phloem conductance in the tree stem. We concluded that prolonged drought affected phloem transport capacity through a change in anatomy and that the slowdown of phloem transport under drought likely resulted from a reduced driving force due to lower hydrostatic pressure between the source and sink organs.

Prevalence of interspecific competition in a mixed poplar/black locust plantation under adverse climate conditions

Key messageIn a mixed poplar/black locust plantation in central France, adverse conditions have led to a prevalence of interspecific competition, resulting in a poorer performance than monocultures.ContextIn mixed tree plantations, the presence of woody N2-fixing species is thought to reduce N needs by fertilization. However, benefits associated to soil nitrogen enrichment have to outweigh the negative effects of interspecific competition. To do so, co-occurring tree species have to be chosen carefully to promote niche sharing between species and reduce competition. Black locust and poplar mixtures therefore seem promising since both species are fast growing and have potentially complementary crown shapes.AimsOur objective was to evaluate the impact of the poplar/black locust mixture on the growth, above- and belowground biomass production, and nitrogen allocation of the two species, as compared to their respective monocultures.MethodsAn experimental plantation mixing poplar and black locust was set up in central France. For five growing seasons, growth, nitrogen allocation, and carbon allocation were monitored for the two species growing either in mixture or in monoculture.ResultsAfter a couple of promising growing seasons, black locust growth and survival slowly declined, mainly in the mixture. At the stand level, biomass production in the mixed plots was nearly 50% below the most productive monoculture (poplar) by age 5xa0years.ConclusionUnder adverse conditions, interspecific competition in the mixture was the preponderant interaction, resulting in higher mortality and lower biomass production than the two monocultures.