Carbon stress causes earlier budbreak in shade-tolerant species and delays it in shade-intolerant species.

PREMISE: Climate change may lead to C stress (negative C balance) in trees. Because nonstructural carbohydrates (NSC) are required during metabolic reactivation in the spring, C stress might delay budbreak timing. This effect is expected to be greater in shade-intolerant than in shade-tolerant species, owing to the faster C economy in the shade-intolerant. METHODS: We experimentally induced C stress in saplings of six temperate tree species that differed in their light requirements by exposing them to either full light or shade from summer to spring, then recorded the date of first budbreak for the individuals. Because the levels of C reserves that represent effective C stress may differ among species, we estimated the degree of C stress by recording survival during the experiment and measuring whole-sapling NSC concentrations after budbreak. RESULTS: Shade reduced NSC concentrations and increased the sugar fraction in the NSC in all species. In the shade, shade-intolerant species had higher mortality and generally lower NSC concentrations than the shade-tolerant species, indicating a trend for more severe C stress in species with faster C economy. In shade-intolerant species, budbreak started earlier and proceeded faster in full light than in shade, but in shade-tolerant species budbreak was delayed in full light. The effects of the light environments on budbreak were not greater in shade-intolerant than in shade-tolerant species. CONCLUSIONS: Our study reveals a correspondence between budbreak responses to light and the light requirements of the species. This finding confirms that C metabolism has a significant role in triggering budbreak and demonstrates that whether C stress accelerates or delays budbreak depends on the species' light requirements.

Inner bark as a crucial tissue for non-structural carbohydrate storage across three tropical woody plant communities.

Non-structural carbohydrates (NSC) are crucial for forest resilience, but little is known regarding the role of bark in NSC storage. However, bark's abundance in woody stems and its large living fraction make it potentially key for NSC storage. We quantified total NSC, soluble sugar (SS) and starch concentrations in the most living region of bark (inner bark, IB), and sapwood of twigs, trunks and roots of 45 woody species from three contrasting tropical climates spanning global extremes of bark diversity and wide phylogenetic diversity. NSC concentrations were similar (total NSC, starch) or higher (SS) in IB than wood, with concentrations co-varying strongly. NSC concentrations varied widely across organs and species within communities and were not significantly affected by climate, leaf habit or the presence of photosynthetic bark. Starch concentration tended to increase with density, but only in wood. IB contributed substantially to NSC storage, accounting for 17-36% of total NSC, 23-47% of SS and 15-33% of starch pools. Further examination of the drivers of variation in IB NSC concentration, and taking into account the substantial contribution of IB to NSC pools, will be crucial to understand the role of storage in plant environmental adaptation.

Carbon allocation to growth and storage depends on elevation provenance in an herbaceous alpine plant of Mediterranean climate.

It is unclear whether the frequently observed increase in non-structural carbohydrates (NSC) in plants exposed to low temperatures or drought reflects a higher sensitivity of growth than photosynthesis in such conditions (i.e. sink limitation), or a prioritization of carbon (C) allocation to storage. Alpine areas in Mediterranean-type climate regions are characterized by precipitation increases and temperature decreases with elevation. Thus, alpine plants with wide elevational ranges in Mediterranean regions may be good models to examine these alternative hypotheses. We evaluated storage and growth during experimental darkness and re-illumination in individuals of the alpine plant Phacelia secunda from three elevations in the Andes of central Chile. We hypothesized that storage is prioritized regarding growth in plants of both low- and high elevations where drought and cold stress are greatest, respectively. We expected that decreases in NSC concentrations during darkness should be minimal and, more importantly, increases in NSC after re-illumination should be higher than increases in biomass. We found that darkness caused a significant decrease in NSC concentrations of both low- and high-elevation plants, but the magnitude of the decrease was lower in the latter. Re-illumination caused higher increase in NSC concentration than in biomass in both low- and high-elevation plants (1.5- and 1.9-fold, respectively). Our study shows that C allocation in Phacelia secunda reflects ecotypic differences among elevation provenances and suggests that low temperature, but not drought, favours C allocation to storage over growth after severe C limitation.

Cluster root-bearing Proteaceae species show a competitive advantage over non-cluster root-bearing species.

BACKGROUND AND AIMS: Cluster roots (CRs) constitute a special root adaptation that enables plants to take up nutrients, especially phosphorus (P), from soils with low nutrient availability, including recent volcanic deposits. It is unclear, however, how CR species interact with non-cluster root-bearing (NCR) species, and how substrates' fertility modulates potential interactions. METHODS: We experimentally assessed the net interaction between CR and NCR species using two substrates of contrasting fertility: nutrient-rich nursery mix and tephra (low P availability). We planted seedlings of two southern South American (SSA) Proteaceae, CR species and two NCR Nothofagus species in pairs (conspecifics and heterospecifics) and as singles. We analysed the effect of seedling neighbours on survival, growth performance (e.g. total biomass and leaf area) and leaf and substrate nutrient concentrations (including manganese, a proxy for P-acquisition efficiency through CR activity) using the relative interaction index. KEY RESULTS: After three growing seasons, we found that (1) Proteaceae species had fewer CRs and lower CR biomass and grew less in the tephra than in the nursery substrate; (2) Nothofagus species did not improve their survival and growth in the presence of Proteaceae species in any substrate; (3) contrary to Nothofagus, Proteaceae species improved their growth more when planted with any neighbour (including conspecifics) than when planted alone, which was accompanied by a significant accretion of leaf P; and (4) the presence of a neighbour increased the final nitrogen and P concentrations in the nursery substrate, regardless of species identity. CONCLUSIONS: CRs provide Proteaceae a competitive advantage over NCR species at the seedling stage, which may have important consequences for species coexistence and community structuring. The investigated SSA Proteaceae, which have not evolved in nutrient-impoverished soils, as have their relatives in south-western Australia and South Africa, improve their growth when cultivated in pairs, especially in nutrient-rich substrates.

Revisiting the relative growth rate hypothesis for gymnosperm and angiosperm species co-occurrence.

PREMISE OF THE STUDY: It is unclear to what extent the co-occurrence of angiosperm and gymnosperm species in some marginal ecosystems is explained by reduced growth in angiosperms due to carbon (C) limitation and by high stress tolerance in gymnosperms associated with lack of vessels and resource conservation. METHODS: We examined growth patterns and traits associated with C balance in four evergreen angiosperm species (including one vesselless species, Drimys winteri) and three gymnosperm tree species of a cold-temperate rainforest in southern Chile. We measured the mean basal area increment for the first 50 (BAI50 ) and the last 10 years (BAI10 ), wood density, leaf lifespan, and nonstructural carbohydrate (NSC) concentrations in different organs. KEY RESULTS: BAI50 was 6-fold higher in angiosperms than in gymnosperms and ca. 4-fold higher in Drimys than in the fastest-growing gymnosperm. BAI10 and aboveground NSC concentrations were significantly higher and leaf lifespan lower in angiosperms than in gymnosperms; these differences, however, were largely driven by the slow growth and low NSC concentrations of the Cupressaceae species (Pilgerodendron uviferum), while the two Podocarpaceae had BAI10 and NSC concentrations similar to angiosperms. In angiosperms, NSC and starch concentrations were generally higher in species with lower BAI10 , indicating no severe C limitation. CONCLUSIONS: The co-occurrence of angiosperms and gymnosperms in cold-temperate rainforests of southern Chile is not explained by growth disadvantages and C limitation in angiosperms. Long leaf longevity, but not lack of vessels, appeared to favor resource conservation and C balance in some gymnosperms (Podocarpaceae).

Herbivore resistance in congeneric and sympatric Nothofagus species is not related to leaf habit.

PREMISE: Two fundamental hypotheses on herbivore resistance and leaf habit are the resource availability hypothesis (RAH) and the carbon-nutrient balance hypothesis (CNBH). The RAH predicts higher constitutive resistance by evergreens, and the CNBH predicts higher induced resistance by deciduous species. Although support for these hypotheses is mixed, they have rarely been examined in congeneric species. METHODS: We compared leaf constitutive and induced resistance (as leaf polyphenol and tannin concentrations, and as damage level in non-choice experiments) and leaf traits associated with herbivory of coexisting Nothofagus species using (1) a defoliation experiment and (2) natural defoliation caused by an outbreak of a common defoliator of Nothofagus species. RESULTS: In the defoliation experiment, polyphenol and tannin concentrations were similar between deciduous and evergreen species; regardless of leaf habit, polyphenols increased in response to defoliation. In the natural defoliation survey, N. pumilio (deciduous) had significantly higher herbivory, lower carbon/nitrogen ratio and leaf mass per area, and higher nitrogen and phosphorus concentrations than N. betuloides (evergreen); N. antarctica (deciduous) had intermediate values. Polyphenol concentrations and herbivore resistance indicated by the non-choice experiment were lower in N. pumilio than in N. antarctica and N. betuloides, which had similar values. CONCLUSIONS: Higher herbivory in N. pumilio was associated with a higher nutritional value and a lower level of leaf carbon-based defenses compared to both the evergreen and the other deciduous species, indicating that herbivore resistance in Nothofagus species cannot be attributed to only leaf habit as predicted by the RAH or CNBH.

Carbon allocation to growth and storage in two evergreen species of contrasting successional status.

PREMISE OF THE STUDY: A prevailing hypothesis in forest succession is that shade-tolerant species grow more slowly than shade-intolerant species, across light conditions, because they prioritize carbon (C) allocation to storage. We examined this hypothesis in a confamilial pair of species, including one of the fastest-growing tree species in the world (Eucalyptus globulus) and a shade-tolerant, slow-growing species (Luma apiculata). METHODS: Seedlings were subjected to one out of four combinations of light (high vs. low) and initial defoliation (90% defoliated vs. nondefoliated) for four months. Growth, C storage concentration in different organs, leaf shedding, and lateral shoot formation were measured at the end of the experiment. KEY RESULTS: Eucalyptus globulus grew faster than L. apiculata in high light, but not in low light. Both species had lower C storage concentration in low than in high light, but similar C storage concentrations in each light condition. Defoliation had no effect on C storage, except in the case of the old leaves of both species, which showed lower C storage levels in response to defoliation. Across treatments, leaf shedding was 96% higher in E. globulus than in L. apiculata while, in contrast, lateral shoot formation was 87% higher in L. apiculata. CONCLUSIONS: In low light, E. globulus prioritized C storage instead of growth, whereas L. apiculata prioritized growth and lateral branching. Our results suggest that shade tolerance depends on efficient light capture rather than C conservation traits.

Extreme defoliation reduces tree growth but not C and N storage in a winter-deciduous species.

BACKGROUND AND AIMS: There is a growing concern about how forests will respond to increased herbivory associated with climate change. Carbon (C) and nitrogen (N) limitation are hypothesized to cause decreasing growth after defoliation, and eventually mortality. This study examines the effects of a natural and massive defoliation by an insect on mature trees' C and N storage, which have rarely been studied together, particularly in winter-deciduous species. METHODS: Survival, growth rate, carbon [C, as non-structural carbohydrate (NSC) concentration] and nitrogen (N) storage, defences (tannins and total polyphenols), and re-foliation traits were examined in naturally defoliated and non-defoliated adult trees of the winter-deciduous temperate species Nothofagus pumilio 1 and 2 years after a massive and complete defoliation caused by the caterpillar of Ormiscodes amphimone (Saturniidae) during summer 2009 in Patagonia. KEY RESULTS: Defoliated trees did not die but grew significantly less than non-defoliated trees for at least 2 years after defoliation. One year after defoliation, defoliated trees had similar NSC and N concentrations in woody tissues, higher polyphenol concentrations and lower re-foliation than non-defoliated trees. In the second year, however, NSC concentrations in branches were significantly higher in defoliated trees while differences in polyphenols and re-foliation disappeared and decreased, respectively. CONCLUSIONS: The significant reduction in growth following defoliation was not caused by insufficient C or N availability, as frequently assumed; instead, it was probably due to growth limitations due to factors other than C or N, or to preventative C allocation to storage. This study shows an integrative approach to evaluating plant growth limitations in response to disturbance, by examining major resources other than C (e.g. N), and other C sinks besides storage and growth (e.g. defences and re-foliation).

Patterns of carbon storage in relation to shade tolerance in southern South American species.

PREMISE OF THE STUDY: Carbon (C) allocation to storage in woody tissues at the expense of growth is thought to promote shade tolerance, yet few studies on the subject examined C storage during maximum growth and considered stand influences. I asked how C storage in different plant tissues relates to shade tolerance in temperate forests with contrasting climates and physiognomies, and whether relationships vary during the growing season. METHODS: In the late spring and late summer, I harvested seedlings of eight species with contrasting light requirements from the understory of a cold rainforest and a Mediterranean forest in Chile. Nonstructural carbohydrate (NSC) concentrations and pools (i.e., biomass x NSC concentration) were determined in leaves, aboveground wood, and roots. The effects of shade tolerance and sampling date on the NSCs were analyzed for each forest and tissue with linear mixed-effects models. KEY RESULTS: In both forests, concentrations of NSC and soluble sugars in woody tissues, as well as fractions of NSC in these tissues, were lower in shade tolerant than in shade intolerant species. For root NSC concentrations, these trends depended on the sampling date: in the late spring the concentrations were similar in shade tolerant and intolerant species, while in the late summer they were lower in shade tolerant species. CONCLUSIONS: Shade tolerance is not linked to C storage in the two studied forests, suggesting that allocation to growth or defenses could be more advantageous for low light persistence. Alternatively, high levels of C storage could be also selected in shade intolerant species to face herbivory or drought.

High foliar nutrient concentrations and resorption efficiency in Embothrium coccineum (Proteaceae) in southern Chile.

PREMISE OF THE STUDY: Southern South American (SA) Proteaceae species growing in volcanic soils have been proposed as potential ecosystem engineers by tapping phosphorus (P) from soil through their cluster roots and shedding nutrient-rich litter to the soil, making it available for other species. We tested whether Embothrium coccineum (Proteaceae) has effectively lower P nutrient resorption efficiency and higher litter P concentrations than co-occurring, non-Proteaceae species. METHODS: In southern Chile, we assessed the P and nitrogen (N) resorption efficiency of senescent leaves and fresh litter of E. coccineum and co-occurring tree species in a soil fertility and moisture gradient (600-3000 mm of annual precipitation) in Patagonia, Chile. We determined P and N concentrations, leaf mass per area (LMA), and ratios of N/P and C/N in mature and senescent leaf cohorts and fresh litter. KEY RESULTS: Embothrium coccineum showed significantly higher P and N resorption efficiency than co-occurring species; in fact, E. coccineum fresh litter had the lowest P-content. While E. coccineum showed significantly lower fresh litter P concentrations than the rest of the species, it showed significantly higher N concentrations. Embothrium coccineum also had lower LMA and similar N/P and C/N ratios when compared with co-occurring tree species. CONCLUSIONS: We found that E. coccineum efficiently mobilized P and, to a lesser extent, N before leaf shedding. We did not find support for the ecosystem engineering hypothesis via shedding P-rich litter. We suggest that southern South American Proteaceae may be taking up other nutrients besides P, probably N, from the young, volcanic soils of this region.

An experimental approach to explain the southern Andes elevational treeline.

UNLABELLED: • PREMISE OF THE STUDY: The growth limitation hypothesis (GLH) is the most accepted mechanistic explanation for treeline formation, although it is still uncertain whether it applies across taxa. The successful establishment of Pinus contorta--an exotic conifer species in the southern hemisphere--above the Nothofagus treeline in New Zealand may suggest a different mechanism. We tested the GLH in Nothofagus pumilio and Pinus contorta by comparing seedling performance and carbon (C) balance in response to low temperatures.• METHODS: At a southern Chilean treeline, we grew seedlings of both species 2 m above ground level, to simulate coupling between temperatures at the meristem and in the air (colder), and at ground level, i.e., decoupling air temperature (relatively milder). We recorded soil and air temperatures as well. After 3 yr, we measured seedling survival and biomass (as a surrogate of growth) and determined nonstructural carbohydrates (NSC).• KEY RESULTS: Nothofagus and Pinus did not differ in survival, which, as a whole, was higher at ground level than at the 2-m height. The root-zone temperature for the growing season was 6.6°C. While biomass and NSC decreased significantly for Nothofagus at the 2-m height compared with ground level (C limitation), these trends were not significant for Pinus• CONCLUSIONS: The treeline for Nothofagus pumilio is located at an isotherm that fully matches global patterns; however, its physiological responses to low temperatures differed from those of other treeline species. Support for C limitation in N. pumilio but not in P. contorta indicates that the physiological mechanism explaining their survival and growth at treeline may be taxon-dependent.

Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients.

BACKGROUND AND AIMS: The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous-evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations. METHODS: Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous-evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra). KEY RESULTS: Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood. CONCLUSIONS: Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.

Soil nitrogen, and not phosphorus, promotes cluster-root formation in a South American Proteaceae, Embothrium coccineum.

PREMISE OF THE STUDY: Cluster roots are a characteristic root adaptation of Proteaceae species. In South African and Australian species, cluster roots promote phosphorus (P) acquisition from poor soils. In a South American Proteaceae species, where cluster roots have been scarcely studied and their function is unknown, we tested whether cluster-root formation is stimulated by low soil nutrition, in particular low P-availability. METHODS: Small and large seedlings (< 6- and > 6-months old, respectively) of Embothrium coccineum and soil were collected across four different sites in Patagonia (Chile). We determined cluster-root number and relative mass, and leaf Pi concentration per mass (Pimass) and per area (Piarea) for each seedling, and tested relationships with Olsen-P (OP), sorbed-P (sP) and total nitrogen (N) using generalized linear mixed-effects models and model selection to assess the relative strength of soil and plant drivers. KEY RESULTS: Best-fit models showed a negative logarithmic relationship between cluster-root number and soil nitrogen (N), and between cluster-root relative mass and both leaf Piarea and soil N, and a positive logarithmic relationship between cluster-root number and leaf Piarea. Cluster-root relative mass was higher in small than in large seedlings. CONCLUSIONS: Contrary to that found in South African and Australian Proteaceae, cluster roots of E. coccineum do not appear to be driven by soil P, but rather by soil N and leaf Piarea. We suggest that cluster roots are a constitutive and functional trait that allows plants to prevail in poor N soils.

Local adaptation to aridity in a widely distributed angiosperm tree species is mediated by seasonal increase of sugars and reduced growth.

Trees in dry climates often have higher concentrations of total non-structural carbohydrates (NSC = starch + soluble sugars) and grow less than conspecifics in more humid climates. This pattern might result from growth being more constrained by aridity than the carbon (C) gain, or reflect local adaptation to aridity, since NSC fuel metabolism and ensure adequate osmoregulation through the supply of soluble sugars (SS), while low growth reduces water and C demands. It has been further proposed that C allocation to storage could come at the expense of growth (i.e., a growth-storage trade-off). We examined whether NSC and growth reflect local adaptation to aridity in Embothrium coccineum (Proteaceae), a species with an exceptionally wide niche. To control for any influence of the phenotypic plasticity on NSC and growth, we collected seeds from dry (500 mm year-1) and moist (> 2500 mm year-1) climates and grew seedlings in a common garden experiment for 3 years. We then compared NSC and SS concentrations and pools (i.e., total contents), and the biomass of seedlings at spring, summer, and fall. Seedlings from the dry climate had significantly lower biomass and similar NSC concentrations and pools than seedlings from moist climate, suggesting that reduced growth in arid environments does not result from a prioritization of C allocation to storage but it confers advantages under aridity (e.g., lower transpiration area). Across organs, starch and NSC decreased similarly in seedlings from both climates from spring onward. However, root and stem SS concentrations increased during the growing season, and these increases were significantly higher in seedlings from the dry climate. The greater SS accumulation in seedlings from the dry climate compared to those from the moist climate demonstrates ecotypic differentiation in the seasonal dynamics of SS, suggesting that SS underlie local adaptation to aridity. (298 words).

Variation of mobile carbon reserves in trees at the alpine treeline ecotone is under environmental control.

In low temperature-adapted plants, including treeline trees, light-saturated photosynthesis is considerably less sensitive to temperature than growth. As a consequence, all plants tested so far show increased nonstructural carbohydrate (NSC) tissue concentrations when exposed to low temperatures. Reduced carbon supply is thus an unlikely cause for low temperature range limits of plants. For altitudinal treeline trees there is, however, a possibility that high NSC genotypes have been selected. Here, we explored this possibility using afforestations with single-provenance conifers along elevational gradients in the Southern Chilean Andes and the Swiss Alps. Tree growth was measured at each of four approximately equidistant elevations at and below the treeline. Additionally, at the same elevations, needle, branch and stem sapwood tissues were collected to determine NSC concentrations. Overall, growth decreased and NSC concentrations increased with elevation. Along with previous empirical and experimental studies, the findings of this study provide no indication of NSC reduction at the treeline; NSC increased in most species (each represented by one common population) towards their upper climatic limit. The disparity between carbon acquisition and structural carbon investment at low temperature (accumulation of NSC) thus does occur even among genotypes not adapted to treeline environments.

Phenology explains different storage remobilization in two congeneric temperate tree species with contrasting leaf habit.

The dependence of trees on carbon and nutrient storage is critical to predicting the forest vulnerability under climate change, but whether evergreen and deciduous species differ in their use and allocation of stored resources during spring phenology is unclear. Using a high temporal resolution, we evaluated the role of spring phenology and shoot growth as determinants of the carbon and nutrient storage dynamics in contrasting leaf habits. We recorded the phenology and shoot elongation and determined the concentrations of total non-structural carbohydrates (NSCs), starch, soluble carbohydrates, nitrogen (N) and phosphorus (P) in buds, expanding shoots and previously formed shoots of two sympatric Nothofagus species with contrasting leaf habit. Species reached similar shoot lengths, though shoot expansion started 35 days earlier and lasted c. 40 days more in the deciduous species. Thus, although the deciduous species had a relatively constant shoot growth rate, the evergreen species experienced a conspicuous growth peak for c. 20 days. In the evergreen species, the greatest decreases in NSC concentrations of previously formed shoots and leaves coincided with the maximum shoot expansion rate and fruit filling, with minimums of 63 and 65% relative to values at bud dormancy, respectively. In contrast, minimum NSC concentrations of the previously formed shoots of the deciduous species were only 73% and occurred prior to the initiation of shoot expansion. Bud N and P concentrations increased during budbreak, whereas previously formed shoots generally did not decrease their nutrient concentrations. Late spring phenology and overlapping of phenophases contributed to the greater dependence on storage of proximal tissues in the studied evergreen compared with deciduous species, suggesting that phenology is a key determinant of the contrasting patterns of storage use in evergreen and deciduous species.

Intraspecific trait variation and covariation in a widespread tree species (Nothofagus pumilio) in southern Chile.

• The focus of the trait-based approach to study community ecology has mostly been on trait comparisons at the interspecific level. Here we quantified intraspecific variation and covariation of leaf mass per area (LMA) and wood density (WD) in monospecific forests of the widespread tree species Nothofagus pumilio to determine its magnitude and whether it is related to environmental conditions and ontogeny. We also discuss probable mechanisms controlling the trait variation found. • We collected leaf and stem woody tissues from 30-50 trees of different ages (ontogeny) from each of four populations at differing elevations (i.e. temperatures) and placed at each of three locations differing in soil moisture. • The total variation in LMA (coefficient of variation (CV) = 21.14%) was twice that of WD (CV = 10.52%). The total variation in traits was never less than 23% when compared with interspecific studies. Differences in elevation (temperature) for the most part explained variation in LMA, while differences in soil moisture and ontogeny explained the variation in WD. Traits covaried similarly in the altitudinal gradient only. • Functional traits of N. pumilio exhibited nonnegligible variation; LMA varied for the most part with temperature, while WD mostly varied with moisture and ontogeny. We demonstrate that environmental variation can cause important trait variation without species turnover.

No evidence of carbon limitation with tree age and height in Nothofagus pumilio under Mediterranean and temperate climate conditions.

BACKGROUND AND AIMS: Trees universally decrease their growth with age. Most explanations for this trend so far support the hypothesis that carbon (C) gain becomes limited with age; though very few studies have directly assessed the relative reductions of C gain and C demand with tree age. It has also been suggested that drought enhances the effect of C gain limitation in trees. Here tests were carried out to determine whether C gain limitation is causing the growth decay with tree age, and whether drought accentuates its effect. METHODS: The balance between C gain and C demand across tree age and height ranges was estimated. For this, the concentration of non-structural carbohydrates (NSCs) in stems and roots of trees of different ages and heights was measured in the deciduous temperate species Nothofagus pumilio. An ontogenetic decrease in NSCs indicates support for C limitation. Furthermore, the importance of drought in altering the C balance with ontogeny was assessed by sampling the same species in Mediterranean and humid climate locations in the southern Andes of Chile. Wood density (WD) and stable carbon isotope ratios (δ(13)C) were also determined to examine drought constraints on C gain. KEY RESULTS: At both locations, it was effectively found that tree growth ultimately decreased with tree age and height. It was found, however, that NSC concentrations did not decrease with tree age or height when WD was considered, suggesting that C limitation is not the ultimate mechanism causing the age/height-related declining tree growth. δ(13)C decreased with tree age/height at the Mediterranean site only; drought effect increased with tree age/height, but this pattern was not mirrored by the levels of NSCs. CONCLUSIONS: The results indicate that concentrations of C storage in N. pumilio trees do not decrease with tree age or height, and that reduced C assimilation due to summer drought does not alter this pattern.

Ontogeny, understorey light interception and simulated carbon gain of juvenile rainforest evergreens differing in shade tolerance.

BACKGROUND AND AIMS: A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried. METHODS: The biomass distribution of juveniles (17-740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain. RESULTS: An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR(d) = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey. CONCLUSIONS: The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.

Low Growth Sensitivity and Fast Replenishment of Non-structural Carbohydrates in a Long-Lived Endangered Conifer After Drought.

There is an ongoing debate on whether a drought induced carbohydrate limitation (source limitation) or a direct effect of water shortage (sink limitation) limit growth under drought. In this study, we investigated the effects of the two driest summers recorded in southern Chile in the last seven decades, on the growth and non-structural carbohydrates (NSC) concentrations of the slow-growing conifer Fitzroya cupressoides. Specifically, we studied the seasonal variation of NSC in saplings and adults one and two years after the occurrence of a 2 year-summer drought at two sites of contrasting precipitation and productivity (mesic-productive vs. rainy-less productive). We also evaluated radial growth before, during and after the drought, and predicted that drought could have reduced growth. If drought caused C source limitation, we expected that NSCs will be lower during the first than the second year after drought. Conversely, similar NSC concentrations between years or higher NSC concentrations in the first year would be supportive of sink limitation. Also, due to the lower biomass of saplings compared with adults, we expected that saplings should experience stronger seasonal NSC remobilization than adults. We confirmed this last expectation. Moreover, we found no significant growth reduction during drought in the rainy site and a slightly significant growth reduction at the mesic site for both saplings and adults. Across organs and in both sites and age classes, NSC, starch, and sugar concentrations were generally higher in the first than in the second year following drought, while NSC seasonal remobilization was generally lower. Higher NSC concentrations along with lower seasonal NSC remobilization during the first post-drought year are supportive of sink limitation. However, as these results were found at both sites while growth decreased slightly and just at the mesic site, limited growth only is unlikely to have caused NSC accumulation. Rather, these results suggest that the post-drought dynamics of carbohydrate storage are partly decoupled from the growth dynamics, and that the rebuild of C reserves after drought may be a priority in this species.