Nutrient content and seasonal fluctuations in the leaf component of coark-oak (Quercus suber L.) litterfall

Nutrient content and seasonality of the leaf component in cork-oak litterfal were studied over a two year period in two cork-oak forest sites differing in biomass and edaphic condition in the north-eastern Iberian peninsula. Fallen senescent leaves compared to young leaves showed higher non-mobile nutrient concentrations and lower mobile nutrient concentrations, specially P, N, K, and Mg. At both sites, seasonal fluctuations affected both leaf production and leaf mineral content. The maximum leaf fall period correspond to the start of the vegetative growth and to the lowest N and P concentration in the falling leaves. The opposite was true for the winter, when litterfal was minimal and N and P content in falling leaves was at a maximum.The comparison between falling leaves and canopy leaves suggests that the analysis of fallen leaves can be a useful measure of N and P nutrition in cork-oak. We found site dependent differences in nutrient content and nutrient remobilization. In comparison with Q. ilex, although litter production was in the same range, nutrient retranslocation was greater for Q. suber.     

The influence of summer and winter stress and water relationships on the distribution of Quercus ilex L.

Several aspects of plant-water relationships and canopy behaviour have been studied over several years in two experimental areas of Quercus ilex L. forests in the northeastern Iberian Peninsula. Water requirements, water status limits necessary for a positive carbon budget and the functional canopy behaviour in the face of abiotic stress were evaluated, in order to determine which factors influence the geographical range of these forests.The results showed that holm oak has a conservative water use, a low cuticular transpiration, a high capacity for osmotic adjustement and xerophytic characteristics in leaf morphology and canopy arrangement. More than 440 mm of annual rainfall are required for these forests to persist. Summer drought and winter cold are thus important abiotic factors limiting the distribution of Quercus ilex. In both cases, drought stress is involved.     

Effects of winter cold stress on photosynthesis and photochemical efficiency of PSII of the Mediterranean <Emphasis Type="Italic">Cistus albidus</Emphasis> L. and <Emphasis Type="Italic">Quercus ilex</Emphasis> L.

This study examined the photosynthetic and growth performances of potted plants of Cistus albidus L. and Quercus ilex L. submitted either to natural Mediterranean winter conditions or to mild greenhouse conditions. Plants grown outdoors exhibited lower light and CO₂-saturated CO₂ assimilation rates (A_sat) and apparent quantum yield (_i) than those indoors. Until mid-winter, C. albidus had higher A_sat than Q. ilex, but differences disappeared after a period of severe cold. Maximal photochemical efficiency of PSII (Fv/Fm) measured predawn was higher in C. albidus than in Q. ilex, and decreased throughout the season in outdoor plants. Fv/Fm also decreased at light saturation (A_sat) in both species. Fv/Fm was correlated with photosynthetic capacity and efficiency (quantum yield), but the resulting regression slopes were different between the two species. At the physiological level, C. albidus seemed to cope better with cold stress than Q. ilex. However, winter stress induced reduction of leaf absorptance, increased leaf mass per area, extensive leaf damage and high plant mortality in C. albidus. This suggests that the high performance of C. albidus leaves is not likely to be maintained for long periods of cold stress, and may therefore depend on continuous leaf replacement. Quercus ilex showed a conservative behaviour, with low net assimilation rates but greater leaf and plant survival than C. albidus.     

Contrasting foliar responses to drought in Quercus ilex and Phillyrea latifolia

Leaf morphology, longevity, and demography were examined in Quercus ilex and Phillyrea latifolia growing in a holm oak forest in Prades mountains (northeast Spain). Four plots (10 × 15 m) of this forest were submitted to an experimental drought during three years (soil moisture was reduced about 15 %). Leaf area, thickness and leaf mass per area ratio (LMA) were measured in sun and shade leaves of both species. Leaf longevity, the mean number of current-year shoots produced per previous-year shoot (Sn/Sn-1), the mean number of current-year leaves per previous-year shoot (Ln/Sn-1), and the percentage of previous-year shoots that developed new ones were measured once a year, just after leaf flushing. LMA and leaf thickness increased since leaf unfolding except in summer periods, when stomatal closure imposed low photosynthetic rates and leaves consumed their reserves. LMA, leaf area, and leaf thickness were higher in Q. ilex than in P. latifolia, but leaf density was higher in the latter species. Drought reduced the leaf thickness and the LMA of both species ca. 2.5 %. Drought also increased leaf shedding up to ca. 20 % in Phillyrea latifolia and decreased it up to ca. 20 % in Q. ilex. In the later species, Sn/Sn-1 decreased by 32 %, Ln/Sn-1 by 41 %, percentage of shoots developed new ones by 26 %, and leaf area by 17 %. Thus the decrease of leaf number and area was stronger in the less drought-resistant Q. ilex, which, under increasingly drier conditions, might lose its current competitive advantage in these Mediterranean holm oak forests.     

Changes in wheat leaf phenolome in response to cold acclimation

A study of wheat (Triticum aestivum L.) leaves phenolome was carried out during cold acclimation of the winter (Claire) and spring (Bounty) varieties using a combination of HPLC–ESI–MS techniques. A total of 40 phenolic and flavonoid compounds were identified, and consisted mainly of two coumarin derivatives, eight simple phenolic derivatives, 10 hydroxycinnamoyl amides and 20 flavonoid derivatives. Identification and quantification of individual compounds were performed using an HPLC system coupled with a photodiode array detector and two different ESI–MS systems, in combination with a multiple reaction monitoring (MRM) technique. The analyses indicated that, although there were no qualitative differences in their profiles, the winter variety exhibited a higher phenolic content compared to the spring variety when both were grown under non-acclimated (control) conditions. Cold acclimation, on the other hand, resulted in a significant differential accumulation of phenolic compounds in both varieties: mostly as luteolin C-glycosides and their O-methyl derivatives in the winter variety (Claire) and a derivative of hydroxycinnamoyl amide in the spring variety (Bounty). These compounds accumulated in relatively large amounts in the apoplastic compartment. The accumulation of the O-methylated derivatives was associated with a marked increase in O-methyltransferase (OMT) activity. In addition, the trimethylated flavone, 3′,4′,5′-trimethyltricetin was identified for the first time in the native extracts of both control and cold-acclimated wheat leaves. The accumulation of a mixture of beneficial flavonoids, such as iso-orientin, vitexin and tricin in cold acclimated wheat leaves, attests for its potential as an inexpensive source of a health-promoting supplement to the human diet.     

Artificially increased ascorbate content affects zeaxanthin formation but not thermal energy dissipation or degradation of antioxidants during cold-induced photooxidative stress in maize leaves

Infiltrating detached maize (Zeamays L.) leaves with L-galactono-1,4-lactone (L-GAL) resulted in a 4-fold increase in the content of leaf ascorbate. Upon exposure to high irradiance (1000 μmol photons m⁻² s⁻¹) at 5 °C, L-GAL leaves de-epoxidized the xanthophyll-cycle pigments faster than the control leaves; the maximal ratio of de-epoxidized xanthophyll-cycle pigments to the whole xanthophyll-cycle pool was the same in both leaf types. The elevated ascorbate content, together with the faster violaxanthin de-epoxidation, did not affect the degree of photoinhibition and the kinetics of the recovery from photoinhibition, assayed by monitoring the maximum quantum efficiency of photosystem II primary photochemistry (F_v/F_m). Under the experimental conditions, the thermal energy dissipation seems to be zeaxanthin-independent since, in contrast to the de-epoxidation, the decrease in the efficiency of excitation-energy capture by open photosystem II reaction centers (F_v′/F_m′) during the high-irradiance treatment at low temperature showed the same kinetic in both leaf types. This was also observed for the recovery of the maximal fluorescence after stress. Furthermore, the elevated ascorbate content did not diminish the degradation of pigments or α-tocopherol when leaves were exposed for up to 24 h to high irradiance at low temperature. Moreover, a higher content of ascorbate appeared to increase the requirement for reduced glutathione. Received: 20 May 1999 / Accepted: 29 October 1999     

Decomposition of litter in sub-alpine forests of Eucalyptus delegatensis,E. pauciflora and E. dives

The rate of decomposition of summer leaf-fall (abscised leaves), winter leaf-fall (containing some green leaves) and mature green (picked) leaves was assessed in sub-alpine forests of E. delegatensis (R. T. Baker), E. pauciflora (Sieb. ex Spreng) and E. dives (Schau.) in the Brindabella Range, Australian Capital Territory, using litter bag and tethered leaf techniques. The relative contribution of leaching, microbial respiration and grazing by invertebrate macrofauna to loss of leaf weight was determined. The effect of leaching and microbial respiration was assessed in terms of weight loss per unit area of leaf (specific leaf weight), while losses due to macro-faunal grazing were assessed by measuring reductions in leaf area. Litter decomposition constants for litter components (leaf, bark, wood) and total litter were determined from long-term records of litterfall and accumulated litter. Weight losses of abscised leaves during the initial 12 months ranged from 25% for E. pauciflora to 39% for E. delegatensis and were almost entirely due to reduction in specific leaf weight. Losses in the weight of leaves falling in winter ranged from 38 to 49%, while green leaves lost 45 - 59%. Approximately 50% of the total weight loss of green leaves was due to a loss in leaf area caused by skeletonization by litter macrofauna. Thus abscised leaves rather than green leaves must be used for measuring litter decomposition rates since abscised leaves constitute most of the litterfall in eucalypt forests. Leaves placed in the field in autumn decomposed slowly during the first summer, while the rate increased during the second winter and summer. Low litter moisture content appears to limit decomposition in the initial summer period in all communities, after which litterfall provides a mulch which reduces the rate of desiccation of lower litter layers. A simple linear regression model relating decomposition rate to the number of days (D) when litter moisture content exceeded 60% ODW accounted for 63-83% of the variation in decomposition of leaves in the field. Inclusion of mean monthly air temperature (T) and the product of D and T (day degrees when litter was wet) in a multiple linear regression increased the variation in decomposition accounted for to 80 – 90%. The rate of weight loss showed a positive linear relationship with the initial concentration of nitrogen (N) or phosphorus (P) in the leaf. These concentrations are an index of the decomposability of leaf substrates (e.g. degree of sclerophylly or lignification). The rate of loss of specific weight was similar for tethered leaves and for leaves enclosed in mesh bags. Measured loss in specific leaf weight after 70 – 90 weeks was less than that predicted using decomposition constants (k).     

Stem xylem features in three Quercus (Fagaceae) species along a climatic gradient in NE Spain

 Stem xylem features in two evergreen Quercus species (Q. coccifera and Q. ilex) and a deciduous one (Q. faginea) were analysed along an Atlantic-Mediterranean climatic gradient in which rainfall and winter cold experience strong variation. Mean maximum vessel diameter, vessel density, vessel element length, xylem transverse sectional area, Huber value (xylem transverse sectional area per leaf area unit), theoretical leaf specific conductivity (estimated hydraulic conductance per leaf area unit) and total leaf area were determined in 3-year-old branches. Q. faginea presented the widest vessels and the highest theoretical leaf specific conductivity while Q. coccifera showed the lowest total leaf area and the highest Huber value. Studied features did not exhibit significant correlations with mean minimum January temperature in any species but did show significant relationships with rainfall. In Q. coccifera, mean maximum vessel diameter, vessel element length and theoretical leaf specific conductivity increased with higher rainfall while vessel density decreased. Mean maximum vessel diameter and total leaf area in Q. ilex increased with precipitation whereas variables of Q. faginea did not show any significant trend. Results suggest that aridity, rather than minimum winter temperature, controls stem xylem responses in the studied evergreen species. Q. faginea traits did not show any response to precipitation, probably because this species develops deep roots, which in turn makes edaphic and topographic factors more important in the control of soil water availability. In response to aridity Q. coccifera only exhibits adjustment at a xylem level by reducing its water transport capacity through a reduction of vessel diameter without changing the amount of xylem tissue or foliage, whereas Q. ilex adjusts its water transport capacity in parallel to the foliage area. Received: 13 January 1997 / Accepted: 8 April 1997     

Comparative photochemical and phenomorphological responses to winter stress of an evergreen (Quercus ilex L.) and a semi-deciduous (Cistus albidus L.) Mediterranean woody species

The impact of winter stress on plants from a Mediterranean area was evaluated through comparison of photosystem II (PS II) efficiencies and phenomorphological characteristics of two Mediterranean woody species – Quercus ilex (evergreen) and Cistus albidus (semi-deciduous). The studies were carried out in NE Spain, at two sites with different mean winter temperatures. The results showed that reductions of the efficiency of PS II may occur in Mediterranean plant communities during winter, and are especially remarkable on colder days. The extent and duration of the decrease in PS II efficiency (photoinhibition) depended not only on the climatic conditions, but also on the site, on the species considered and on the position of leaves in the plant canopy. Increased photoinhibition at the colder site was only clear for C. albidus. Nevertheless, the efficiencies of PS II were always higher in C. albidus than in Q. ilex. Phenological patterns, morphology and leaf inclination may protect C. albidus leaves from potentially photoinhibitory conditions in winter. Morphological and structural photoprotection is apparently not so well developed in Q. ilex, which probably relies more on increased protection at the photochemical level of its long-lived leaves. As has been reported in relation to summer-drought stress, Q. ilex has possibly developed a strategy of tolerance to photoinhibition, whereas C. albidus relies preferentially on avoidance features.     

Foliar flavonoids of eastern North AmericanVitis (Vitaceae) north of Mexico

Twenty-one flavonoid glycosides were isolated from the leaves of 22 North AmericanVitis L. taxa, representing two subgenera and five series. Three chemical groups were evident: one producing flavonols, flavones, and C-glycosylflavones, a second producing flavonols and flavones, and a third producing only flavonols. These three chemical groups did not correspond to any of the subgeneric groupings based on morphology. However, flavonoid distributions within series in each subgenus correlate well with morphological data. Parallel flavonoid evolution within each series is thought to account for this lack of subgeneric and interserial flavonoid distinction. The flavonoid data indicate that seriesCordifoliae of subgenusVitis, particularlyV. vulpina L., is the most closely related group to subgenusMuscadinia (Planch.)Rehder, and represents an evolutionary link between the two subgenera.     

Predicting the Recovery of <Emphasis Type="Italic">Pinus halepensis</Emphasis> and <Emphasis Type="Italic">Quercus ilex</Emphasis> Forests after a Large Wildfire in Northeastern Spain

Quercus ilex and Pinus halepensis are two of the most common tree species of the western Mediterranean basin. Both species regenerate reliably after fire: P. halepensis colonizes recently disturbed areas by effective seedling recruitment, while Q. ilex resprouts vigorously after disturbances. For this reason, the natural regeneration of these species after fire should ensure the re-establishment of a forest similar to that which existed before the fire. This study analyzes with a simple simulation model whether or not the relative abundance of monospecific and mixed forests of these species in the landscape is altered by fire. We also analyze the topographic factors and the forest structure before the fire that determine the changes in forest composition after fire. This study has been carried out in a large fire that occurred in NE Spain. Overall, 33% of plots changed to another community type, but this probability of change varied considerably among community types before the fire. Monospecific forests of P. halepensis or Q. ilex had a high probability of remaining in their original composition after the fire, whereas the resilience of mixed forests of these two species was quite low. Mixed forests changed for the most part to monospecific P. halepensis or Q. ilex forests. Analysis of several factors determining these changes indicated that only elevation as a significant topographical variable. The effect of fire was to increase the altitudinal differentiation between the two species. P. halepensis forests that changed to mixed or Q. ilex forests were those of highest elevation, while the mixed and Q. ilex plots that changed to P. halepensis forests were those located at the lowest elevations. Concerning structural variables before fire, density of Q. ilex trees before the fire showed a much greater effect than P. halepensis density in determining the post-fire community. Finally, burn severity also influenced the changes observed. For both P. halepensis and Q. ilex forests, plots that changed to another forest type were mainly those that burned more severely. In the case of mixed forests, even low fire severities involved high probabilities of change to monospecific forests.     

Macroinvertebrate community patterns in relation to leaf-associated periphyton under contrasting light and nutrient conditions in headwater streams

1. Temperate headwater streams traditionally have been considered heterotrophic and brown food web dominated with little primary production. Recent work, however, suggests algae on leaves in these streams may play a greater role than previously thought through interactions with microbial decomposers like fungi. Algae also may be important for macroinvertebrates colonizing leaves in streams. Algae are a more nutritious food resource for shredders than fungi and bacteria and provide a food resource for non-shredder macroinvertebrates.
2. In a field experiment, we manipulated light in three low-nutrient and three high-nutrient streams using leaf bags filled with red maple leaves in winter and spring. After four weeks we measured algal and fungal biomass, leaf stoichiometry, and macroinvertebrate abundance and biomass associated with the leaf bags. We also identified the macroinvertebrate community and examined differences in functional feeding guilds and taxa under ambient- and shaded-light treatments and low- and high-nutrient concentrations in relation to measured leaf characteristics.
3. Algal biomass on leaves was greatest in high-nutrient streams and ambient-light treatments in both seasons. Fungal biomass on leaves was greatest in high-nutrient streams and showed a moderate marginally significant positive correlation with algae during the winter. Leaf C:N was negatively correlated to algae in winter and fungi in both seasons, while leaf N:P and C:P were negatively correlated to fungi in winter and algae in spring. Interactions between fungi and algae on leaves and the nutritional importance of each for macroinvertebrates likely change across seasons, potentially impacting macroinvertebrate community composition.
4. Macroinvertebrate diversity did not differ, but biomass was significantly greater in shaded-light treatments during spring. Abundance was highest in the high-nutrient ambient-light conditions in both seasons, corresponding to greatest algal biomass. Functional feeding guild biomass and abundance were related to different leaf characteristics by season and guild. Higher algal biomass was an important factor for colonization of certain macroinvertebrates (e.g., Ephemerella (Ephemeroptera: Ephemerellidae) and Stenonema (Ephemeroptera: Heptageniidae)), while others were more abundant under shaded treatments with lower algal biomass (e.g., Tipula (Diptera: Tipulidae)), indicating taxa-specific responses.
5. Leaf-associated algae may be an important factor mediating macroinvertebrate communities associated with leaves in temperate headwater streams. Our results demonstrate that green and brown food webs intersect within leaf packs, and they cannot be easily disentangled. We therefore should consider both autochthonous and allochthonous resources within headwater streams when examining their communities or developing water management strategies.

     

Heteroblasty in Arabidopsis thaliana (L.) Heynh

 Heteroblasty in Arabidopsis thaliana was analyzed in a variety of plants with mutations in leaf morphology using a tissue-specific β-glucuronidase gene marker. Some mutants exhibited their mutant phenotypes specifically in foliage leaves. The phenotypes associated with the foliage-leaf-specific mutations were also found to be induced ectopically in cotyledons in the presence of the lec1 mutation. Moreover, the features of an emf1lec1 double mutant showed that cotyledons can be partially converted into carpelloids. When heteroblastic traits were examined in foliage leaves in the presence of certain mutations or natural deviations by histochemical analysis of the expression of the tissue-specific marker gene, it was found that ectopic expression of the developmental program for the first foliage leaves in lec1 cotyledons seemed to affect the heteroblastic features of the first set of foliage leaves, while foliage leaves beyond the third position appeared normal. Similarly, in wild-type plants, discrepancies in heteroblastic features, relative to standard features, of foliage leaves at early positions seemed to be eliminated in foliage leaves at later positions. These results suggest that heteroblasty in foliage leaves might be affected in part by the heteroblastic stage of the preceding foliage leaves but is finally controlled autonomously at each leaf position. Received: 9 July 1999 / Accepted: 17 August 1999     

Expression of low molecular weight heat-shock proteins and total antioxidant activity in the Mediterranean tree Quercus ilex L. in relation to seasonal and diurnal changes in physiological parameters

The total antioxidant activity (TAA) and the accumulation of small heat shock proteins (sHsps) were analysed under field conditions in Quercus ilex with regard to organ ontogeny and the physiological state of the plant. The results point toward the participation of sHsps and an increase of TAA in the general adaptation syndrome (GAS) of woody Mediterranean evergreens. In leaves and stems, there was a definite TAA seasonal pattern but no effect from diurnal variations or from the current stage of organ ontogeny. TAA was about twice as large in summer as in spring and winter and the hydrophilic antioxidant content was about 16 times greater than that of lipophilic antioxidants. The accumulation of sHsps in leaves also showed a seasonal pattern, but no effect from diurnal variations or from leaf ontogeny. In summer days, the sHsps content remained invariable even during the daylight hours in which the leaves were physiologically recovered. However, the accumulation of sHsps in stems did vary in relation to organ ontogeny. Old stems had a high accumulation of sHsps throughout the year, whereas in young stems, accumulation of sHsps was detected only in summer. This is probably due to a higher quantity of lignified and suberized tissues in the older stems. Using two‐dimensional immunodetection for leaves and stems, two sets of Hs protein species (17 and 10 kDa regions) were observed. In stems, there was an increase in 10 kDa proteins from winter to summer. These results are discussed and the possible role of the two types of polypeptides in the face of environmental and endogenous oxidative stress are considered.     

Leaf Anatomy,Inclination, and Gas Exchange Relationships in Evergreen Sclerophqldous and Drought Semideciduous Shrub Species

There are significant differences in leaf life-span among evergreen sclerophyllous species and drought semideciduous species growing in the Mediterranean maquis. Cistus incamus, which has a leaf life-span of four-eight months, was characterised by the highest net photosynthetic rates (P _N), while Quercus ilex and Phillyrea latifolia, which maintain their leaves two-three and two-four years, respectively, had a lower P _N. The longer leaf life-span of the two evergreen sclerophyllous species may be justified to cover the high production costs of leaf protective structures such as cuticle, hairs, and sclereids: cuticle and hairs screen radiation penetrating into the more sensitive tissues, and sclereids have a light-guiding function. Q. ilex and P. latifolia have the highest leaf mass/area ratio (LMA = 209 g m^-2) and a mesophyll leaf density (2065 cells per mm² of leaf cross section area) about two times higher than C. incanus. In the typical evergreen sclerophyllous species the steepest leaf inclination ( = 56°) reduces 42 % of radiation absorption, resulting in a reduced physiological stress at leaf level, particularly in summer. C. incanus, because of its low leaf life-span, requires a lower leaf investment in leaf protective structures. It exhibits a drastic reduction of winter leaves just before summer drought, replacing them with smaller folded leaves. The lower leaf inclination ( = 44°) and the lower LMA (119 g m^-2) of C. incanus complement photosynthetic performance. Water use efficiency (WUE) showed the same trend in Q. ilex, P. latifolia, and C. incanus, decreasing 60 % from spring to summer, due to the combined effects of decreased CO₂ uptake and increased transpirational water loss.     

不同色彩珙桐叶片和苞片解剖结构及色素含量比较研究

以生长于同一生境下的粉红珙桐(粉红色叶片、苞片)与普通珙桐(绿色叶片、白色苞片)为试材,对比两种色彩珙桐叶片/苞片解剖结构和色素含量的差异,以揭示珙桐色彩转变的规律。结果显示：(1)两种珙桐叶片均属于异面叶类型,栅栏组织由一层长柱形细胞整齐排列而成,海绵组织排列疏松,部分粉红叶片的上表皮细胞向外凸起,绿叶无此现象;粉红叶片的总厚度及其表皮角质层、栅栏组织和海绵组织厚度都高于绿叶,而表皮较薄。(2)两种珙桐苞片均无栅栏组织和海绵组织的分化,粉红苞片上表皮细胞明显隆起,上表皮角质层增厚,而下表皮变薄。(3)粉红叶片的类黄酮、花色苷含量分别是绿色叶片的1.52倍、3.67倍,两者的光合色素含量无显著差异,但粉红叶片的叶绿素a/b值比绿色叶低很多;粉红苞片花色苷含量显著高于白色苞片,而两者类黄酮含量差异不大。研究表明,花色苷是珙桐叶片和苞片色彩转红的直接因素,类黄酮有助于叶片呈红色;粉红珙桐叶片/苞片的解剖结构发生了一定变化,对光能的利用效率更高,对阴湿环境的适应性增强。     

Variation in tissue element concentrations in Quercus ilex L. over a range of different soils

In order to study the variability in nutrient concentrations in four tissues of Q. ilex in relation to soil properties, we selected fifteen stands in both Quercus ilex forests and Q. ilex-Pinus halepensis mixed forests. These stands had developed on soils derived from eight different parent materials. Three soil groups were differentiated according to their chemical properties: calcareous soils, siliceous soils, and volcanic soils. Across sites, nutrient concentrations were generally less variable in current-year tissues than in older tissues. Nitrogen and potassium showed the lowest variability among sites, their concentrations in current-year leaves ranging from 1.17% to 1.39% for N and from 0.53% to 0.68% for K. There were few statistically significant correlations between tissue element concentrations, the most frequent being the antagonistic relationship between calcium and magnesium. Nitrogen concentration in current-year leaves was negatively correlated with soil chemical fertility (nitrogen, phosphorus and potassium). This may reflect a nutritional imbalance between nitrogen and other nutrients, some of which may be more limiting than nitrogen to Q. ilex growth in Catalonia forests. Negative correlations were also found between plant magnesium and soil calcium, and positive correlations between plant calcium and soil calcium.     

The seasonal dynamics of leaf resin,nitrogen, and herbivore damage in Eriodictyon californicum and their parallels in Diplacus aurantiacus

Summary The chaparral shrub Eriodictyon californicum produces a flavonoid leaf resin with a chemically similar composition to that previously reported for the sympatric shrub Diplacus aurantiacus. We determined the phenology, resin content, and nitrogen content of Eriodictyon leaves and the leaf area lost to herbivores. Nitrogen content and resin content were both negatively correlated with leaf age at each sampling date, but nitrogen decreased during the growing season while resin increased. The fraction of leaf area lost during the growing season averaged less than 7% and was highest on the oldest leaves. The seasonal pattern of resin production in Eriodictyon corresponds to that in Diplacus, indicating that the similar environments of Eriodyctyon and Diplacus have led to convergent leaf resins. This convergence in these two plants has implications for chemicals of similar function in other chaparral shrubs.     

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.     

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.