Shade Tolerance (shade + tolerance)

Distribution by Scientific Domains


Selected Abstracts


Allometry and Shade Tolerance in Pole-Sized Trees of Two Contrasting Dipterocarp Species in Sabah, Malaysia,

BIOTROPICA, Issue 3 2006
Martin G. Barker
ABSTRACT We compared the allometry of two contrasting late-successional dipterocarp species to test whether a monolayer (shade-tolerant),multilayer (shade-intolerant) model applies to pole-sized trees. Crown traits of the more shade-tolerant species (Vatica micrantha) did not conform to either of the familiar monolayer or multilayer models for pole-sized trees, but instead were consistent with a "persistent multilayer" model. Species differences in crown traits may be influenced more by future rather than present light environments. [source]


Xylem root and shoot hydraulics is linked to life history type in chaparral seedlings

FUNCTIONAL ECOLOGY, Issue 1 2010
Robert B. Pratt
Summary 1.,Shrubs in fire prone chaparral communities have evolved different life history types in response to fire. A key to understanding the evolution of life history type differences is to understand how physiological traits are linked to differences in life history type. Vascular adaptations are important for delivering an efficient and stable water supply to evergreen chaparral shrub leaves. This study tested for a link between vascular physiology and life history type in chaparral shrubs. 2.,Chaparral shrub species along the south-western coast of North America survive wildfire by three different life histories. Non-sprouters are killed by fire and re-establish exclusively through germination of fire-stimulated seeds, facultative sprouters re-establish by a combination of vegetative sprouting and fire-stimulated seeds, and obligate sprouters re-establish exclusively by vegetative sprouting because their seeds do not survive fire. Non-sprouters and facultative sprouters establish seedlings in the open canopy post fire environment, whereas obligate sprouters establish seedlings in the shady understory of the mature chaparral canopy. 3.,Seedlings of nine species (Rhamnaceae) representing three each of the different life history types were grown in deep containers in a common garden under treatments of sun and shade. Hydraulic conductance was measured using a high-pressure flow meter for all organs, and a vacuum technique was used to measure conductance of fine and woody roots. We predicted that non-sprouters would exhibit greater hydraulic efficiency than the sprouting species, and that facultative sprouters would be more efficient than the shade tolerant obligate sprouters. 4.,Non-sprouters had the greatest hydraulic conductance per unit leaf and sapwood area at the whole seedling level, whereas facultative and obligate sprouters were not different. Comparing hydraulic conductance across major organs (from fine roots to leaves) showed that the hydraulic system was well coordinated. At the whole seedling level, the root system was more of a bottleneck than the shoot system. This pattern was consistent with high resistance extraxylary pathways in roots and differences in root architecture. 5.,The greater hydraulic efficiency of the non-sprouter life history type is attributed to its post-fire pioneering habit and may partially explain the relatively high speciation in the non-sprouters. Lower hydraulic efficiency is associated with a sprouting life history and greater shade tolerance. The seedling root systems represent a hydraulic bottleneck that may place roots under especially intense selection. [source]


Allocation of above-ground growth is related to light in temperate deciduous saplings

FUNCTIONAL ECOLOGY, Issue 4 2003
D. A. King
Summary 1Allocational shifts in response to light may be an important factor in allowing plants to survive in shade, while increasing their extension rates and competitive ability in sun. To investigate this response, the allocation of above-ground growth between leaves, branches and stems was studied in saplings of Acer pensylvanicum L. and Castenea dentata (Marsh.) Borkh. in the Appalachian mountains of western Virginia, USA. Measurements of current leaf biomass, current and past year leaf numbers and the growth ring widths of branches and stem were used to estimate biomass partitioning for saplings growing in locations ranging from forest understorey to large openings. 2Both species showed higher leaf area per unit leaf biomass (SLA) and higher allocation of above-ground growth to leaves in shade than in sun. 3There were no differences between species in the slopes of the relationships of allocation and SLA vs estimated irradiance, but SLA was significantly greater in A. pensylvanicum than in C. dentata at a given light level. Hence, somewhat lower production per unit leaf area is required to maintain the canopy in A. pensylvanicum, consistent with foresters' ratings of greater shade tolerance for this species. 4Greater foliar allocation in shade than sun has also been observed in broad-leaved evergreen saplings, but generally not in seedlings. This difference is probably related to differences in size and age between seedlings and saplings. Young seedlings typically show exponential growth with no immediate foliar losses, while shaded saplings lie closer to the steady state where new leaves replace old ones with little additional stem growth. 5Thus trees shift their allocation patterns in an acclimatory fashion, depending on their size and light environment, with the costs of replacing senesced leaves becoming of consequence as juveniles age. [source]


Do shade-tolerant tropical tree seedlings depend longer on seed reserves?

FUNCTIONAL ECOLOGY, Issue 4 2002
Functional growth analysis of three Bignoniaceae species
Summary 1.,A functional growth analysis was used to determine the duration of strict dependency on seed reserves for energy and nitrogen in three woody Bignoniaceae species (Tabebuia rosea DC., Challichlamys latifolia K. Schum. and Pithecoctenium crucigerum A. Gentry) which differed in cotyledon function (photosynthetic, semi-photosynthetic and storage) and shade tolerance (probability of seedling establishment and survival in the understorey). 2.,Seedlings were raised from seeds in sand culture under combinations of three nitrogen levels (daily supply of nutrient solution containing 100, 10 and 0% of 2·6 mm N) and two irradiances (27 and 1% full sun). Time course of biomass, non-cotyledonous biomass and leaf area for 40 days post-germination were compared to identify when the external availability of nitrogen or light began to affect seedling growth. 3.,Seedlings of all species became dependent on external energy supply earlier than they did on nitrogen supply. In all species seed nitrogen was sufficient to support positive seedling growth for 40 days in shade, but not in sun. 4.,Tabebuia rosea with photosynthetic cotyledons responded to light availability earlier than more shade-tolerant species with storage cotyledons. Challichlamys latifolia, the most shade-tolerant species, had the highest nitrogen concentration in seeds and was the last to respond to external nitrogen availability. Thus seedlings of the most shade-tolerant species depended on seed reserves for the longest period for both energy and nitrogen. 5.,Relative growth rate after seedlings initiated autotrophic growth was in a trade-off relationship with seedling survivorship in the understorey across the three species. Tabebuia rosea, the least shade-tolerant species, had the highest positive net carbon balance in sun and shade. 6.,Functional morphology of cotyledons and concentration of seed nitrogen deserve as much attention as seed size as correlates of contrasting seedling regeneration strategies. [source]


Trade-offs in low-light CO2 exchange: a component of variation in shade tolerance among cold temperate tree seedlings

FUNCTIONAL ECOLOGY, Issue 2 2000
M. B. Walters
Abstract 1.,Does enhanced whole-plant CO2 exchange in moderately low to high light occur at the cost of greater CO2 loss rates at very-low light levels? We examined this question for first-year seedlings of intolerant Populus tremuloides and Betula papyrifera, intermediate Betula alleghaniensis, and tolerant Ostrya virginiana and Acer saccharum grown in moderately low (7·3% of open-sky) and low (2·8%) light. We predicted that, compared with shade-tolerant species, intolerant species would have characteristics leading to greater whole-plant CO2 exchange rates in moderately low to high light levels, and to higher CO2 loss rates at very-low light levels. 2.,Compared with shade-tolerant A. saccharum, less-tolerant species grown in both light treatments had greater mass-based photosynthetic rates, leaf, stem and root respiration rates, leaf mass:plant mass ratios and leaf area:leaf mass ratios, and similar whole-plant light compensation points and leaf-based quantum yields. 3.,Whole-plant CO2 exchange responses to light (0·3,600 µmol quanta m,2 s,1) indicated that intolerant species had more positive CO2 exchange rates at all but very-low light (< 15 µmol quanta m,2 s,1). In contrast, although tolerant A. saccharum had a net CO2 exchange disadvantage at light > 15 µmol quanta m,2 s,1, its lower respiration resulted in lower CO2 losses than other species at light < 15 µmol quanta m,2 s,1. 4.,Growth scaled closely with whole-plant CO2 exchange characteristics and especially with integrated whole-plant photosynthesis (i.e. leaf mass ratio × in situ leaf photosynthesis). In contrast, growth scaled poorly with leaf-level quantum yield, light compensation point, and light-saturated photosynthetic rate. 5.,Collectively these patterns indicated that: (a) no species was able to both minimize CO2 loss at very-low light (i.e. < 15 µmol quanta m,2 s,1) and maximize CO2 gain at higher light (i.e. > 15 µmol quanta m,2 s,1), because whole-plant respiration rates were positively associated with whole-plant photosynthesis at higher light; (b) shade-intolerant species possess traits that maximize whole-plant CO2 exchange (and thus growth) in moderately low to high light levels, but these traits may lead to long-term growth and survival disadvantages in very-low light (< 2·8%) owing, in part, to high respiration. In contrast, shade-tolerant species may minimize CO2 losses in very-low light at the expense of maximizing CO2 gain potential at higher light levels, but to the possible benefit of long-term survival in low light. [source]


The effect of tree height and light availability on photosynthetic leaf traits of four neotropical species differing in shade tolerance

FUNCTIONAL ECOLOGY, Issue 1 2000
T. Rijkers
Abstract 1.,Light-saturated rate of photosynthesis (Amax), nitrogen (N), chlorophyll (Chl) content and leaf mass per unit area (LMA) were measured in leaves of trees of different heights along a natural light gradient in a French Guiana rain forest. The following four species, arranged in order from most shade-tolerant to pioneer, were studied: Duguetia surinamensis, Vouacapoua americana, Dicorynia guianensis and Goupia glabra. Light availability of trees was estimated using hemispherical photography. 2.,The pioneer species Goupia had the lowest LMA and leaf N on both an area and mass basis, whereas Duguetia had the highest values. In general, leaf variables of Vouacapoua and Dicorynia tended to be intermediates. Because Amax/area was similar among species, Goupia showed both a much higher light-saturated photosynthetic nitrogen-use efficiency (PNUEmax) and Amax/mass. Leaves of Vouacapoua demonstrated the greatest plasticity in Amax/area, particularly in small saplings. 3.,A distinction could be made between the effect of tree height and light availability on the structural, i.e. LMA, and photosynthetic leaf characteristics of all four species. The direction and magnitude of the variation in variables were similar among species. 4.,LMA was the key variable that mainly determined variation in the other leaf variables along tree height and light availability gradients, with the exception of changes in chlorophyll concentration. Amax/area, N/area, LMA and stomatal conductance to water vapour (gs) increased, whereas Chl/mass decreased, with both increasing tree height and canopy openness. Amax/mass, PNUEmax and Amax/Chl increased with increasing openness only. N/mass and Chl/area were independent of tree height and openness, except for small saplings of Goupia which had a much lower Chl/area. [source]


Size-dependence of growth and mortality influence the shade tolerance of trees in a lowland temperate rain forest

JOURNAL OF ECOLOGY, Issue 4 2009
Georges Kunstler
Summary 1A trade-off between growth in high-light and survival in low-light of species is often proposed as a key mechanism underpinning the dynamics of trees in forest communities. Yet, growth and survival are known to depend on plant size and few studies have analysed how this trade-off can vary between juvenile life stages and the potential consequences of the trade-off for the differences in regeneration rate between species in mixed forests. 2We quantified growth and mortality for two different juvenile life stages , seedlings and saplings , of seven tree species common in temperate rain forests in New Zealand using data from field studies. We found strong evidence that the ranking of species for survival in shade and growth in full light was affected by size. There was a trade-off between seedling survival in low light and sapling height growth in high light, but no trade-offs were observed when considering other combinations of life stages (seedling growth vs. seedling survival, seedling growth vs. sapling survival, or sapling growth vs. sapling survival). 3We ran simulations with an individual-based forest dynamics model , SORTIE/NZ , to explore how the trade-off drives the differences in tree species regeneration success in gaps vs. under closed forest conditions. These simulations indicate that because species' ranks in shade tolerance varied with life stage, regeneration success was not predicted from knowledge of tree performance at a single life stage. For instance, high-light sapling growth was a strong determinant of regeneration success in forest gaps, but seedling growth was also influential. Under closed forest, regeneration success was primarily limited by low-light mortality at the seedling stage, but seedling growth and sapling survival were also influential. 4Synthesis. Growth-survival trade-offs can be strongly affected by the size of the individual analysed, resulting in completely different rankings of the shade tolerance of species across different juvenile life stages. Performance of both seedlings and saplings influenced regeneration success, highlighting the need to consider growth-survival trade-offs and the shade-tolerance strategies of tree species over a large range of juvenile sizes. [source]


The stratification theory for plant coexistence promoted by one-sided competition

JOURNAL OF ECOLOGY, Issue 3 2009
Takashi Kohyama
Summary 1It is an essential feature of plants that leaves at higher levels have better access to light than those at lower levels. Thus, larger plants generally enjoy greater success in competing for light than smaller ones. We analyse the effect of such size-asymmetry, or one-sided competition, on the successful coexistence of plant species, using an analytically tractable model for stratified populations, in which a plant in the same layer exhibits the same crowding effect as any other, irrespective of species. 2A two-layer population that is reproductive in upper layer and juvenile in lower layer has a uniquely stable (plant-size-weighted) equilibrium density, as long as its fecundity is sufficient to compensate for its mortality rate. We also calculate a unique threshold lower-layer density of this layered population when there is no upper-layer plant. This threshold lower-layer density is larger than the weighted equilibrium density with upper layer, except for the case of perfect two-sided competition. 3A two-layer species can stably coexist with a one-layer, understorey species as a result of one-sided, but not two-sided competition. The coexistence condition is that the equilibrium density of the one-layer species lies between the threshold lower-layer density and the equilibrium density of the two-layer species. For an understorey species to coexist successfully with a two-layer species, any advantage in demographic performance, most prominently in a sufficiently high fecundity per plant must offset the disadvantage of living in dark conditions. 4Results from a model of multi-layer populations suggest that several species differing in terms of maximum layer and fecundity can coexist under conditions of one-sided competition. We demonstrate an example of the stable coexistence of eight species. The inter-specific trade-offs predicted by the model correspond to patterns observed in a rain forest. 5Synthesis. We propose a stratification theory that explains the generation and maintenance of the successful coexistence of plant species. Under the condition of one-sided competition, a canopy population that takes advantage of escaping from understorey competition shows an ability to invade an understorey with a density higher than its own equilibrium density, and which offers opportunities for an understorey population with high fecundity and/or shade tolerance to coexist. The predicted coexistence of species that share maximum canopy height is most pronounced for trees of tropical rain forests. [source]


Above- versus below-ground competitive effects and responses of a guild of temperate tree species

JOURNAL OF ECOLOGY, Issue 1 2009
K. David Coates
Summary 1The neutral theory debate has highlighted the scarcity of robust empirical estimates of the magnitude of competitive effects and responses within guilds of co-occurring tree species. Our analysis quantifies the relative magnitude of all possible pairwise competitive interactions within a guild of nine co-occurring tree species in temperate forests of northern, interior British Columbia, and explicitly partitions the competitive effects of neighbours into the effects of shading versus the residual effects of ,crowding', assumed to reflect below-ground competition. 2Models that treated neighbours as equivalent in their competitive effects were the most parsimonious for the five species with the smallest sample sizes. For the remaining species (samples sizes of > 150 individuals), the best models estimated separate competition coefficients for all nine species of neighbours. We take this as evidence that species do indeed differ in their competitive effects, but that there can be a minimum sample size required to discriminate between them. 3There was a strong size-dependency in potential growth. Six species showed an optimal growth at a size between 5 and 20 cm diameter. Potential growth declined moderately to strongly as diameter increased. Sensitivity to crowding varied as a function of tree size for five of the nine species; however, this response was not consistent by tree species. 4The magnitude of reduction in growth due to crowding was greater on average than the reduction in growth due to shading, except for the two least shade tolerant conifers. Sensitivity to shading among the conifer species was correlated with their shade tolerance. 5The per capita effects of crowding by different species of neighbours varied widely. A large number of the estimated pairwise per capita competition coefficients were very low. The relative magnitude of the strength of intra- versus interspecific competition also varied widely among the tree species. 6Synthesis. Model selection techniques effectively separated above- and below-ground competition in complex forests, and allowed us to assess differences among species in competitive effects and responses. While below-ground effects were strong, they were due to proximity of neighbours from a very specific (and small) subset of strong competitors within the guild. Response to crowding varied with tree size but the nature of the relationship varied widely among the species. [source]


Carbohydrate storage enhances seedling shade and stress tolerance in a neotropical forest

JOURNAL OF ECOLOGY, Issue 2 2007
JONATHAN A. MYERS
Summary 1To survive in forest understoreys, seedlings must depend on carbohydrate reserves when they experience negative carbon balance imposed by occasional light reduction and tissue loss to herbivores and diseases. We present the first experimental evidence in support of this hypothesis, using seven woody neotropical species. 2We transplanted seedlings that had recently expanded their first photosynthetic cotyledon or leaf to the forest understorey (1% of full sun) and quantified initial biomass and total non-structural carbohydrate (TNC) in stems, roots and storage cotyledons. We then randomly assigned seedlings to control and two stress treatments: light reduction (0.08% of full sun for 8 weeks) and complete defoliation. 3First-year survival of control seedlings, a comparative measure of shade tolerance, differed widely among species. The two stress treatments reduced survival and relative growth rates (RGR) of all species. Shade-tolerant species were little impacted by the stress treatments, whereas the two least shade-tolerant species experienced 100% mortality. 4In all treatments, 8-week and first-year survival was positively correlated with initial TNC pool size in stems and roots. By contrast, survival was generally not correlated with initial TNC concentration in any organ, TNC pools in cotyledons, seed mass or seedling biomass. 5TNC in stems and roots, but not in cotyledons, decreased in response to light reduction and defoliation over 8 weeks. Leaf area recovery of defoliated seedlings was positively correlated with initial TNC pools in stems and roots. 6First-year survival in each treatment was negatively correlated with 0,8 week RGR of control seedlings, suggesting higher stress tolerance of species with inherently slow growth rates in shade. RGR of control seedlings from 0 to 8 weeks was negatively correlated with initial TNC pools, but not concentrations, in stems and roots. After 8 weeks, RGR was positive for all species, without clear relationships with survival or TNC. 7We conclude that carbohydrate storage in stems and roots enhances long-term survival in shade by enabling seedlings to cope with periods of biotic and abiotic stress. Carbohydrate storage is a key functional trait that can explain species differences in growth and survival that lead to species coexistence through niche assembly processes and life-history trade-offs. [source]


The millennial dynamics of a boreal forest stand from buried trees

JOURNAL OF ECOLOGY, Issue 3 2004
DOMINIQUE ARSENEAULT
Summary 1We reconstructed the dynamics of a black spruce (Picea mariana) and jack pine (Pinus banksiana) forest stand in northern Québec using a continuous, 5200-year-long sequence of stem remains buried in adjacent peatland. Simulations of recruitment of such remains provided guidelines for inferring past ecosystem structure and composition at the stand scale. 2Compared with the late Holocene (4650,0 cal. year BP (CYBP)), the mid Holocene (5200,4650 CYBP) period was characterized by faster tree growth, larger stems and higher stem density, indicating higher forest productivity in association with a milder climate. 3The presence of stem remains of both species from 17 out of 20 contiguous 250-year time intervals suggests that the spruce-pine stand exhibited high compositional stability, with both species regenerating after fire from canopy-stored seed banks. 4Relative species abundance closely followed the duration of past fire intervals deduced from the number of tree rings in buried conifers. Time periods of long (4650,3950, 3400,1850 and 250,0 CYBP) and short fire intervals (4950,4650, 3950,3400 and 1850,250 CYBP) were associated, respectively, with decreasing and increasing pine abundance, probably reflecting faster juvenile growth, lower shade tolerance, earlier sexual maturity and shorter longevity in jack pine compared with black spruce. 5We conclude that both climate change and climate-induced fire disturbance have been driving long-term ecosystem dynamics. Our field evidence supports the idea that interactions between disturbances and the life-history traits of species modulate the impact of climate change at the scale of forest stands. At the same time, disturbances may result in long-term stability of disturbance-adapted ecosystems. [source]


The influence of environmental factors, the pollen : ovule ratio and seed bank persistence on molecular evolutionary rates in plants

JOURNAL OF EVOLUTIONARY BIOLOGY, Issue 1 2006
C.-A. WHITTLE
Abstract One of the main goals of molecular evolutionary biology is to determine the factors that influence the evolutionary rate of selectively neutral DNA, but much remains unknown, especially for plants. Key factors that could alter the mutation rate include environmental tolerances (because they reflect a plants vulnerability to changes in habitat), the pollen : ovule ratio (as it is associated with the number of mitotic divisions) and seed longevity (because this influences the number of generations per unit time in plants). This is the first study to demonstrate that seed bank persistence and drought tolerance are positively associated with molecular evolutionary rates in plants and that pollen : ovule ratio, shade tolerance and salinity tolerance have no detectable relationship. The implications of the findings to our understanding of the impact of environmental agents, the number of cell divisions and cell aging on neutral DNA sequence evolution are discussed. [source]


Comparative demography of three coexisting Acer species in gaps and under closed canopy

JOURNAL OF VEGETATION SCIENCE, Issue 1 2008
H. Tanaka
Abstract Questions: 1. Is there a trade-off between gap dependency and shade tolerance in each of the life-history stages of three closely related, coexisting species, Acer amoenum (Aa), A. mono (Am) and A. rufinerve (Ar)? 2. If not, what differences in life-history traits contribute to the coexistence of these non-pioneer species? Location: Ogawa Forest Reserve, a remnant (98 ha), species-rich, temperate deciduous forest in central Japan (36°56' N, 140°35' E, 600 - 660 m a.s.l.). Methods: We estimated the demographic parameters (survival, growth rate and fecundity) by stage of each species growing in gaps and under closed canopy through observations of a 6-ha permanent plot over 12 years. Population dynamics were analysed with stage-based matrix models including gap dynamics. Results: All of the species showed high seedling and sapling survival rates under closed canopies. However, demographic parameters for each growth stage in gaps and under closed canopies revealed inter-specific differences and ontogenetic shifts. The trade-off between survival in the shade and growth in gaps was detected only at the small sapling stage (height < 30 cm), and Ar had the highest growth rate both in the shade and in the gaps at most life stages. Conclusions: Inter-specific differences and ontogenetic shifts in light requirements with life-form differences may contribute to the coexistence of the Acer species in old-growth forests, with Aa considered a long-lived sub-canopy tree, Am a long-lived canopy tree, and Ar a short-lived,,gap-phase' sub-canopy tree. [source]


Plant traits as predictors of woody species dominance in climax forest communities

JOURNAL OF VEGETATION SCIENCE, Issue 3 2001
Fumito Koike
Satake et al. (1989) Abstract. The dominance of a given tree or shrub species in a particular forest community may be determined by many ecological traits of the target species, as well as those of the surrounding species as its potential competitors. The present study was conducted to evaluate the possibility of predicting community status (species composition and dominance) on the basis of traits of local flora using statistical methods, and to visualize the mathematical function which determines species dominance. A general linear model and logistic regression were used for the statistical analysis. Dependent variables were designated as dominance and presence/absence of species in climax forest, with independent variables as vegetative and reproductive traits. Subalpine, cool-temperate, warm-temperate and subtropical climax rain forests in East Asia were studied. Quantitative prediction of climax community status could readily be made based on easily measured traits of local flora. Species composition and 74.6% of the total variance of species dominance were predicted based on two traits; maximum height and shade tolerance. Through application of this method, the capacity of an alien species to invade a climax forest community could possibly be predicted prior to introduction of the alien species. [source]


Antioxidant and Pigment Composition during Autumnal Leaf Senescence in Woody Deciduous Species Differing in their Ecological Traits

PLANT BIOLOGY, Issue 5 2003
J. I. García-Plazaola
Abstract: Photoprotection mechanisms have been studied during autumnal senescence in sun and shade leaves of woody plants with different ecological characteristics and senescence patterns. Three of them belonging to the same family, Betulaceae: the shade-intolerant and early successional species (Betula alba L.), the shade-tolerant and late successional species (Corylus avellana L.), and an N-fixing tree with low N resorption efficiency (Alnus glutinosa L.). The other two species: a shade-intolerant (Populus tremula L.) and a shade-tolerant (Cornus sanguinea L.), were chosen because of their ability to accumulate anthocyanins during autumnal leaf senescence. The study of plants with different ecological strategies allowed us to establish general trends in photoprotection mechanisms during autumnal senescence, when nutrient remobilisation occurs, but also during whole leaf ontogeny. We have not found a clear relationship between shade tolerance and the level of photoprotection; the main difference between both groups of species being the presence of ,-carotene in shade leaves of shade-tolerant species. Preceding autumn, nitrogen resorption started in mid-summer and occurred in parallel with a slight and continuous ascorbate, chlorophyll and carotenoid degradation. However, the ascorbate pool remained highly reduced and lipid oxidation did not increase at this time. Contrasting with ascorbate, ,-tocopherol accumulated progressively in all species. Only during the last stages of senescence was chlorophyll preferentially degraded with respect to carotenoids, leading to the yellowing of leaves, except in A. glutinosa in which a large retention of chlorophyll and N took place. Senescing leaves were characterised, except in C. sanguinea, by a relative increase in the proportion of de-epoxidised xanthophylls: zeaxanthin, antheraxanthin and lutein. The light-induced accumulation of anthocyanins in C. sanguinea could play an additional protective role, compensating for the low retention of de-epoxidised xanthophylls. These different strategies among deciduous species are consistent with a role for photoprotective compounds in enhancing nitrogen remobilization and storage for the next growing season. [source]