Preferential Feeding and Occupation of Sunlit Leaves Favors Defense Response and Development in the Flea Beetle,Altica brevicollis coryletorum – A Pest of Corylus avellana

The monophagous beetle, Altica brevicollis coryletorum, is a major leaf pest of Corylus avellana (common hazel). In contrast to majority of the other studied species of shrubs, sunlit leaves are grazed to a much greater extent than shaded leaves. Since the observation of a link between leaf irradiance level and A. brevicollis feeding is unique, we hypothesized that feeding preference of this beetle species is related to the speed needed to escape threats i.e. faster jumping. We also hypothesized that sunlit leaves are more nutritious and easier to consume than the leaves of shaded shrubs. Results indicated that beetle mass was greater in beetles occupying sunlit leaves, which is consistent with our second hypothesis. The study also confirmed under laboratory conditions, that larvae, pupae and beetles that were fed full-light (100% of full light) leaves were significantly heavier than those fed with shaded leaves (15% of full light). In the high irradiance conditions (higher temperature) duration of larval development is also reduced. Further results indicated that neither the concentration of soluble phenols, leaf toughness, or the number of trichomes could explain the insect’s preference for sunlit leaves. Notably, measurements of jump length of beetles of this species, both in the field and under laboratory conditions, indicated that the defense pattern related to jumping was associated with light conditions. The jump length of beetles in the sun was significantly higher than in the shade. Additionally, in laboratory tests, beetle defense (jumping) was more strongly affected by temperature (15, 25, or 35°C for 24h) than by leaf type. The effect of sunlit, higher nutrient leaves (greater level of non-structural carbohydrates) on defense (jumping) appears to be indirect, having a positive effect on insect mass in all developmental stages.     

Carotenoid composition in sun and shade leaves of plants with different life forms

The carotenoid composition of sun leaves of nine species of annual crop plants (some with several varieties) was compared with sun and shade leaves of several other groups of plants, among those sun and shade leaves of several species of perennial shrubs and vines and deep-shade leaves of seven rainforest species. All sun leaves contained considerably greater amounts of the components of the xanthophyll cycle violaxanthin, antheraxanthin and zeaxanthin as well as of β-carotene than the shade leaves, as had previously been reported for a variety of other species by Thayer & Björkman (Photosynthesis Research, 1990, 23, 331–343). Therefore, high light specifically stimulated β,β-carotenoid synthesis. The sun leaves of these crop species did not contain α-carotene which was, however, present in large amounts in all shade leaves and in smaller amounts in sun leaves of three of the four species of perennial shrubs and vines. There was no difference in neoxanthin content on a chlorophyll basis between sun and shade leaves, and there was no consistent general difference in the lutein content between all sun and all shade leaves. The zeaxanthin (and antheraxanthin) content at peak irradiance and the xanthophyll cycle pool size were compared for sun leaves from the different groups of plants with different life forms and different metabolic activities. When growing in full sunlight the annual crop species and a perennial mesophyte had high rates of photosynthesis whereas the perennial shrubs and vines had relatively low photosynthesis rates. More zeaxanthin (and antheraxanthin) were accumulated at noon in full sunlight in those species with the lower photosynthesis rates. However, it was not such that those species also possessed the larger pools of violaxanthin + antheraxanthin + zeaxanthin. Instead, the xanthophyll cycle pools of sun leaves of the annual crop species and the perennial mesophyte were not smaller, and were even possibly larger, than those of sun leaves of the perennial shrubs and vines with low photosynthesis rates. This was so in spite of the fact that the crop species experienced much lesser degrees of excessive light at full sun than the shrubs and vines. Thus, many of the crop species converted only about 30–50% of their xanthophyll cycle pool to zeaxanthin at noon, whereas the shrubs and vines typically converted more than 80% of their pool into zeaxanthin. The crop species also had larger pools of β-carotene than the shrubs and vines but smaller pools of lutein than the majority of the latter species.     

Light as a regulator of structural and chemical leaf defenses against insects in two Prunus species

Light is a key factor influencing competition between species, and the mechanisms by which trees overcome insect outbreaks can be associated with alternation of the leaves structure, which then prevent or promotes their susceptibility to herbivores. It was predicted that leaf tissue anatomy would likely be different in sun and shade leaves, with a gradual decline of leaves resistance coupled with reduction of accessible light. We quantified anatomical patterns and the distribution of defence compounds (phenols, total tannins, catechol tannins) within heavily grazed leaves of Prunus padus, native in Europe and Prunus serotina, an invasive to Central Europe. Both species were strongly attacked by folivorous insects when shrubs grew in the shade. In the sun, however only P. padus leaves were grazed, but P. serotina leaves were almost unaffected. We identified that anatomical characteristics are not linked to different P. padus and P. serotina leaf vulnerability to insects. Furthermore, the staining of defence compounds of P. serotina leaves grown in full sun revealed that the palisade mesophyll cells had a higher content of phenolic compounds and catechol tannins. Thus, our results indicate that a specific distribution of defence compounds, but not the anatomical relationships between palisade and spongy mesophyll, may be beneficial for P. serotina growth outside its natural range. The identified pattern of defence compounds distribution is linked to a lower susceptibility of P. serotina leaves to herbivores, and is associated with its invasiveness. This likely reflects that P. serotina is a stronger competitor than P. padus, especially at high sunlit sites i.e. gaps in the forest.     

Leaf diffusion resistance pattern in an oak-hornbeam forest

Four tree, five shrub, and ten herbaceous species growing naturally in an oak-hornbeam forest were used for simultaneous study of the leaf diffusive resistances in the course of several summer days. Absolute minima of the stomatal resistance in the sun tree, the shrub, and the herbaceous species leaves were 1.7 to 6.2 s cm^-1, 6.1 to 10.8 s cm^-1, and 4.8 to 9.7 (17.3 inConvallaria majalis leaves) s cm^-1, respectively. Minimum daily leaf resistances in the course of a day were noted earlier in the morning in sun leaves of large trees than in shade leaves of other species. Stomata were fully opened later in the morning and they began to close sooner in the afternoon in usual shade leaves of the plants in the interior of the forest canopy than those in sun leaves in active surfaces of the canopy (tops of tree crowns). The relatively large differences in leaf resistances found among investigated species may be explained by differences in leaf anatomy (stomata frequency and size) and in ambient leaf or plant environment caused by leaf (plant) position in different vertical layers.     

Nonstructural carbohydrate allocation following different frequencies of simulated browsing in three semi-arid shrubs

Nonstructural carbohydrate allocation patterns in response to different frequencies of simulated browsing (leaf and twig removal) were studied in the following semi-arid shrubs: Osteospermum sinuatum, a dwarf deciduous shrub, Pteronia pallens, a dwarf evergreen shrub, and Ruschia spinosa, a dwarf leaf-succulent shrub. Simulated browsing at all frequencies resulted in the elevation, or had no effect, on total nonstructural carbohydrate (TNC) concentrations of O. sinuatum plant parts, and resulted in the decrease in TNC concentrations of R. spinosa plant parts. The responses of P. pallens were intermediate with elevations as well as declines in TNC concentrations of plant parts measured in response to various clipping frequencies. At the low frequency of simulated browsing (every 26 weeks) elevations in plant TNC content were measured in the two non-succulent shrubs O. sinuatum and P. pallens. It was concluded that the overcompensation with respect to TNC accumulation observed in the two non-succulent species represents one of the ways in which excess photosynthate is utilized by browsed shrubs with a limited regiowth potential. Simulated browsing was the least detrimental with respect to biomass production to the non-succulent O. sinuatum and P. pallens, and most injurious to the leaf-succulent shrub, R. spinosa. The observed TNC allocation patterns could not adequately explain the variation among species in the production of new growth and it was concluded that some factor(s) other than the carbon resource was limiting regrowth.     

SPRING GROWTH OF SHOOTS AND ROOTS IN SHRUBS OF AN ALASKAN MUSKEG

Early spring shoot and fine-root development of four evergreen and three deciduous shrub species were analyzed in a subarctic muskeg at Fairbanks, Alaska. The overwintered foliage of the evergreen shrubs regreened earlier than new leaves developed on the deciduous species. Likewise, the evergreen shrubs produced new fine roots earlier than the deciduous species. The total nonstructural carbohydrate (TNC) concentration did not decline in the evergreen shrub Ledum palustre during the spring development. This contrasted with the deciduous shrub Betula glandulosa, where a significant TNC reduction in stem tissue coincided with bud break and fine-root growth flush.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

More manganese accumulates in maple sun leaves than in shade leaves

Nuclear magnetic resonance and neutron activation analysis were used to measure manganese concentrations in leaves of Acer platanoides. Mn was found to accumulate in both the vacuoles and the chloroplasts, with more Mn (per unit area) in sun leaves than in shade leaves. No Mn was lost at senescence. Different seasonal patterns of Mn accumulation were found in sun and shade leaves. The quantity of chloroplast reserve Mn (bound to the outer surface of thylakoid membranes) increased rapidly in sun leaves from bud-break through midsummer, and then remained approximately constant through senescence. In shade leaves, however, the quantity of reserve Mn increased slowly, and at approximately a constant rate throughout the growing season.     

Carbon allocation to defense, storage, and growth in seedlings of two temperate broad-leaved tree species

Optimal carbon allocation to growth, defense, or storage is a critical trait in determining the shade tolerance of tree species. Thus, examining interspecific differences in carbon allocation patterns is useful when evaluating niche partitioning in forest communities. We hypothesized that shade-tolerant species allocate more carbon to defense and storage and less to growth compared to shade-intolerant species. In gaps and forest understory, we measured relative growth rates (RGR), carbon-based defensive compounds (condensed tannin, total phenolics), and storage compounds (total non-structural carbohydrate; TNC) in seedlings of two tree species differing in shade tolerance. RGR was greater in the shade-intolerant species, Castanea crenata, than in the shade-tolerant species, Quercus mongolica var. grosseserrata, in gaps, but did not differ between the species in the forest understory. In contrast, concentrations of condensed tannin and total phenolics were greater in Quercus than in Castanea at both sites. TNC pool sizes did not differ between the species. Condensed tannin concentrations increased with increasing growth rate of structural biomass (GRstr) in Quercus but not in Castanea. TNC pool sizes increased with increasing GRstr in both species, but the rate of increase did not differ between the species. Accordingly, the amount of condensed tannin against TNC pool sizes was usually higher in Quercus than in Castanea. Hence, Quercus preferentially invested more carbon in defense than in storage. Such a large allocation of carbon to defense would be advantageous for a shade-tolerant species, allowing Quercus to persist in the forest understory where damage from herbivores and pathogens is costly. In contrast, the shade-intolerant Castanea preferentially invested more carbon in growth rather than defense (and similar amounts in storage as Quercus), ensuring establishment success in gaps, where severe competition occurs for light among neighboring plants. These contrasting carbon allocation patterns are closely associated with strategies for persistence in these species’ respective habitats.     

Interaction effects of root-zone salinity and solar irradiance on the physiology and biochemistry of Olea europaea

Root-zone salinity stress and high solar irradiance concomitantly occurs in the Mediterranean basin, where Olea europaea is the dominating fruit-tree crop-species. Although the effect of each individual stressor on plant performance has been widely investigated, much less is known on the interaction effects of salinity stress and solar irradiance on the physiology and biochemistry of olive plants. Here we analyzed how changes in root-zone NaCl concentration and sunlight radiation affect relevant physiological and biochemical features in olive cv. Cipressino. Two-year-old plants were supplied with 0 or 125 mM NaCl and exposed to 15% (shade) or 100% sunlight (sun) over a 5-week period, starting from July 10th, 2005. Measurements were conducted of (i) gas exchange and plant growth, (ii) the concentrations of cations and chloride, (iii) the concentrations of soluble carbohydrates, violaxanthin-cycle pigments and polyphenols, and (iv) the protein oxidation and the lipid peroxidation in the leaves. Salt-induced reductions in gas exchange performance and plant growth were greater at the sun than at the shade site, mostly due to light-induced changes in leaf water relations and vapour pressure deficit (vpd), rather than in the concentration of potentially toxic ions. Light-induced increases in leaf Na⁺ and Cl^? concentrations were countered by parallel enhancements in the concentrations of K⁺ and Ca²⁺. Sun leaves had sharply greater concentrations of mannitol and xanthophylls, irrespective of root-zone salinity. The amount of “newly assimilate carbon” allocated to polyphenols, especially to flavonoids, increased in response to salinity stress and high sunlight. Remarkably, the protein oxidation was greater in shade than in sun leaves of well-watered plants, and increased more at the shade than at the sun site because of high salinity. We suggest that heat-stress (on average maximum T exceeded 33 °C for 50% of the experimental period), which acted in concert with salinity stress and sunlight irradiance in determining plant responses in our experiment, was responsible for leaf oxidative damage in plants growing under contrasting solar radiation. Indeed, sun leaves of salt-stressed plants were equipped with an extraordinary-rich arsenal of antioxidant compounds, distributed in different cell compartments, i.e., mannitol, zeaxanthin and flavonoids, which likely countered effectively the oxidative damage driven by heat-stress, a clear example of cross-tolerance.     

Leaf plasticity in response to light of three evergreen species of the Mediterranean maquis

Morphological, anatomical, biochemical and physiological traits of sun and shade leaves of adult Quercus ilex, Phillyrea latifolia and Pistacia lentiscus shrub species co-occurring in the Mediterranean maquis at Castelporziano (Latium) were studied. Fully expanded sun leaves had 47% (mean of the three species) greater leaf mass area (LMA) and 31% lower specific leaf area (SLA) than shade leaves. Palisade parenchyma thickness contributed on an average 42% to the total leaf thickness, spongy layer 43%, upper epidermal cells 5%, and upper cuticle thickness 3%. Stomatal size was greater in sun (25.5 μm) than in shade leaves (23.6 μm). Total chlorophyll content per fresh mass was 71% greater in shade than in sun leaves, and nitrogen content was the highest in sun (13.7 mg g⁻¹) than in shade leaves (11.8 mg g⁻¹). Difference of net photosynthetic rates (P _N) between sun and shade leaves was 97% (mean of the three species). The plasticity index (sensu Valladares et al., New Phytol 148:79–91, 2000a) was the highest for physiological leaf traits (0.86) than for morphological, anatomical and biochemical ones. Q. ilex had the highest plasticity index of morphological, anatomical and physiological leaf traits (0.37, 0.28 and 0.71, respectively) that might explain its wider ecological distribution. The higher leaf plasticity of Q. ilex might be advantageous in response to varying environmental conditions, including global change.     

Influence of native and alien Prunus species and light conditions on performance of the leaf beetle Gonioctena quinquepunctata

The polyphagous beetle Gonioctena quinquepunctata Fabricius (Coleoptera: Chrysomelidae) is a serious leaf pest of the native European bird cherry, Prunus padus L., and the invasive alien black cherry, Prunus serotina Ehrh. (Rosaceae). In the shade, leaf damage is extensive in both species, whereas in full light, it is extensive in P. padus, but very low in P. serotina. We determined the influence of Prunus species and light conditions on differences in performance of both sexes of this folivore. In a laboratory experiment in which larvae were fed with leaves of a single species grown under particular light conditions, we measured larval, pupal, and adult mass, efficiency of conversion of ingested food (ECI), duration of development, total food eaten, and relative growth rate. In the field, we observed differences in beetle mass on shrubs of the two species growing under various light conditions. From the field observations, we hypothesised that leaves of the invasive P. serotina are not an equally good food source as leaves of P. padus for G. quinquepunctata, and the preference of these beetles for shaded shrubs is most favourable for their growth and development. Under laboratory conditions, we found that the beetle growth rate was not affected significantly by Prunus species or light conditions, despite the significant effect of light condition on the structure and chemical composition of Prunus seedlings. The lower ECI value for larvae feeding on sunlit leaves was compensated for by their higher level of consumption. In the field, adult insect mass was higher on P. padus than on P. serotina, and higher on sunlit shrubs of both species than on shaded ones. Under natural conditions, the mass of adult insects is probably also affected by other factors, such as predators and competition among folivores.     

Seasonal dynamics of non-structural carbohydrates in two species of mediterranean sub-shrubs with different leaf phenology

The seasonal dynamics of non-structural carbohydrates in the woody organs of two co-existing mediterranean sub-shrubs were analyzed. The two species show different leaf phenology during summer: Linum suffruticosum, maintains many of its green leaves, while Lepidium subulatum sheds most of its leaves. These different leaf phenologies are related to different strategies with regard to summer stress. The maintenance of leaves in Linum is related to its stress tolerance while Lepidium avoids stress by shedding its leaves. The main objectives were to: (1) determine the differences in the seasonal dynamics of non-structural carbohydrates among the main woody organs of both species; (2) verify if differences in the leaf phenology, and hence in the strategy with regard to summer drought, lead to different seasonal patterns of carbohydrate storage and use between the two species; (3) compare the seasonal dynamics of carbohydrates of the two studied sub-shrubs with those of mediterranean trees and shrubs previously reported in the literature. The concentration of soluble sugars (SS), starch and total non-structural carbohydrates (TNC) were assessed monthly, over 17 months, in the main roots, stems and the transition zone between root and shoot systems of both species. Starch storage capacity and SS, starch and TNC pools were calculated. The seasonal pattern of carbohydrate accumulation was similar among the woody organs analyzed, but it differed with those reported for mediterranean trees and shrubs. The two species showed different pools and seasonal patterns of non-structural carbohydrate concentrations in its woody organ, which corresponded to their different extent of leaf shedding. The stress-avoider Lepidium accumulated starch during spring shoot growth as a carbon store for summer respiration and had low pools of SS, whereas the stress-tolerant Linum increased SS during summer drought to maintain photosynthetic activity during summer and had low starch pools and storage capacity. However, irrespective of their different leaf shedding patterns, both species had a similar relative variation of their TNC concentration, which contrasts with previous results on deciduous and evergreen woody species.     

Dynamic acclimation to sunlight in an alpine plant,Soldanella alpina L.

In the French Alps, Soldanella alpina (S. alpina) grow under shade and sun conditions during the vegetation period. This species was investigated as a model for the dynamic acclimation of shade leaves to the sun under natural alpine conditions, in terms of photosynthesis and leaf anatomy. Photosynthetic activity in sun leaves was only slightly higher than in shade leaves. The leaf thickness, the stomatal density and the epidermal flavonoid content were markedly higher, and the chlorophyll/flavonoid ratio was significantly lower in sun than in shade leaves. Sun leaves also had a more oxidised plastoquinone pool, their PSII efficiency in light was higher and their non-photochemical quenching (NPQ) capacity was higher than that of shade leaves. Shade-sun transferred leaves increased their leaf thickness, stomatal density and epidermal flavonoid content, while their photosynthetic activity and chlorophyll/flavonoid ratio declined compared to shade leaves. Parameters indicating protection against high light and oxidative stress, such as NPQ and ascorbate peroxidase, increased in shade-sun transferred leaves and leaf mortality increased. We conclude that the dynamic acclimation of S. alpina leaves to high light under alpine conditions mainly concerns anatomical features and epidermal flavonoid acclimation, as well as an increase in antioxidative protection. However, this increase is not large enough to prevent damage under stress conditions and to replace damaged leaves.     

Characterization of pigment apparatus in winter-green and summer-green leaves of a shade-tolerant plant <Emphasis Type="Italic">Ajuga reptans</Emphasis>

Comparative study was performed to assess the content and proportions of photosynthetic pigments and the violaxanthin cycle (VXC) activity in winter-green and summer-green leaves of bugleweed (Ajuga reptans L.) plants grown in shaded (photosynthetically active radiation, PAR 150 μmol/(m² s)) and sunny (PAR 1200 μmol/(m² s)) habitats in the Botanic Garden of Jagiellonian University (Krakow, Poland). In overwintered and newly formed leaves of shade plants, the content of green and yellow pigments was two times higher than in leaves of sun plants. The shade plants were distinguished by accumulation of β-carotene, while lutein was predominant in leaves of sun plants. Under the action of strong light (2000 μmol/(m²s)), the level of violaxanthin deepoxidation in winter-green leaves of shade and sun plants increased five- to sixfold, whereas it changed insignificantly in summer-green leaves of shade plants. It is concluded that, in a shadetolerant species A. reptans, the photosynthetic apparatus of winter-green leaves in sun and shade plants and of summer-green leaves in sun plants is protected against excess insolation by high activity of VXC. The carotenoids of summer-green leaves in shade plants are supposed to function mainly as light-harvesting pigments.     

How do Mediterranean shrub species cope with shade? Ecophysiological response to different light intensities

• Under natural conditions, light exposure for Mediterranean shrubs can be highly variable, especially during cloudy days or under a canopy, and can interfere with other environmental factors such as temperature and water availability.
• With the aim of decoupling the effect of radiation and temperature from water availability, we conducted an experiment where two perennial and three summer semi‐deciduous shrub species were subjected to different levels of irradiation. In order to follow plant responses to light exposure, we measured gas exchange, photosystem II photochemical efficiency, photosynthetic pigments and leaf mass area in spring and summer.
• Results showed that all study species presented a plastic response to different light conditions, and that light‐related traits varied in a coordinated manner. Summer semi‐deciduous species exhibited a more opportunistic response, with higher photosynthesis rates in full sun, but under shade conditions, the two strategies presented similar assimilation rates. Stomatal conductance did not show such a drastic response as photosynthetsis, being related to changes in WUE. Daily cycles of F_v/F_m revealed a slight photoinhibitory response during summer, mainly in perennial species. In all cases photosynthetic pigments adjusted to the radiation level; leaves had lower chlorophyll content, higher pool of xanthophylls and higher proportion of the de‐epoxydaded state of xanthophylls under sun conditions. Lutein content increased in relation to the xanthophyll pool under shade conditions.
• Our results evidenced that radiation is an important driving factor controlling morphological and physiological status of Mediterranean shrub species, independently of water availability. Summer semi‐deciduous species exhibit a set of traits with higher response variability, maximising their photosynthetic assimilation under different sun conditions.

     

Is Shade Beneficial for Mediterranean Shrubs Experiencing Periods of Extreme Drought and Late-winter Frosts?

Background and Aims

Plants are naturally exposed to multiple, frequently interactive stress factors, most of which are becoming more severe due to global change. Established plants have been reported to facilitate the establishment of juvenile plants, but net effects of plant–plant interactions are difficult to assess due to complex interactions among environmental factors. An investigation was carried out in order to determine how two dominant evergreen shrubs (Quercus ilex and Arctostaphylos uva-ursi) co-occurring in continental, Mediterranean habitats respond to multiple abiotic stresses and whether the shaded understorey conditions ameliorate the negative effects of drought and winter frosts on the physiology of leaves.

Methods

Microclimate and ecophysiology of sun and shade plants were studied at a continental plateau in central Spain during 2004–2005, with 2005 being one of the driest and hottest years on record; several late-winter frosts also occurred in 2005.

Key Results

Daytime air temperature and vapour pressure deficit were lower in the shade than in the sun, but soil moisture was also lower in the shade during the spring and summer of 2005, and night-time temperatures were higher in the shade. Water potential, photochemical efficiency, light-saturated photosynthesis, stomatal conductance and leaf ¹³C composition differed between sun and shade individuals throughout the seasons, but differences were species specific. Shade was beneficial for leaf-level physiology in Q. ilex during winter, detrimental during spring for both species, and of little consequence in summer.

Conclusions

The results suggest that beneficial effects of shade can be eclipsed by reduced soil moisture during dry years, which are expected to be more frequent in the most likely climate change scenarios for the Mediterranean region.Key words: Frost, climate change, shade, drought, plant–plant interactions, Quercus ilex, Arctostaphylos uva-ursi, soil moisture, facilitation     

Photosynthetic responses to light in seedlings of selected Amazonian and Australian rainforest tree species

Summary Seedlings of the Caesalpinoids Hymenaea courbaril, H. parvifolia and Copaifera venezuelana, emergent trees of Amazonian rainforest canopies, and of the Araucarian conifers Agathis microstachya and A. robusta, important elements in tropical Australian rainforests, were grown at 6% (shade) and 100% full sunlight (sun) in glasshouses. All species produced more leaves in full sunlight than in shade and leaves of sun plants contained more nitrogen and less chlorophyll per unit leaf area, and had a higher specific leaf weight than leaves of shade plants. The photosynthetic response curves as a function of photon flux density for leaves of shade-grown seedlings showed lower compensation points, higher quantum yields and lower respiration rates per unit leaf area than those of sun-grown seedlings. However, except for A. robusta, photosynthetic acclimation between sun and shade was not observed; the light saturated rates of assimilation were not significantly different. Intercellular CO₂ partial pressure was similar in leaves of sun and shade-grown plants, and assimilation was limited more by intrinsic mesophyll factors than by stomata. Comparison of assimilation as a function of intercellular CO₂ partial pressure in sun- and shade-grown Agathis spp. showed a higher initial slope in leaves of sun plants, which was correlated with higher leaf nitrogen content. Assimilation was reduced at high transpiration rates and substantial photoinhibition was observed when seedlings were transferred from shade to sun. However, after transfer, newly formed leaves in A. robusta showed the same light responses as leaves of sun-grown seedlings. These observations on the limited potential for acclimation to high light in leaves of seedlings of rainforest trees are discussed in relation to regeneration following formation of gaps in the canopy.     

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

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 α‐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, α‐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.     

Photochemical efficiency of adult and young leaves of the neotropical understory shrub Psychotria limonensis (Rubiaceae) in response to changes in the light environment

We explored the short-term adjustment in photochemical efficiency (Fv/Fm) in adult and young leaves of the understory neotropical shrub Psychotria limonensis Krause (Rubiaceae) in response to rapid changes in the light environment. Leaves were collected from 20 individual plants growing under sun and shade conditions on Gigante Peninsula, Barro Colorado Natural Monument (Republic of Panama), during the wet season of 1996. Leaves were distributed in four sequences of light treatments (AB leaves were expanded under sun and were transferred to shade, BA leaves experienced the opposite transfer, and the controls AA and BB leaves that were expanded and maintained under sun or shade conditions). Adult and young leaves did not differ in overall photochemical efficiency. Instead, differences were found among light environments, for which leaves transferred from shade to sun showed the lowest Fv/Fm ratios. There was no relationship between photochemical efficiency and leaf temperature. In P. limonensis, understory plants are susceptible of photoinhibition independently of the leaf ontogenetic stage. The approach utilized in this experiment allowed the rapid exploration of this capacity, and could be applied to poorly studied understory species.