Structural and functional variability within the canopyand its relevance for carbon gain and stress avoidance

The functional variability in leaf angle distribution within the canopy was analysed with respect to regulation of light interception and photoprotection. Leaf orientation strongly determined the maximum photochemical efficiency of PSII (F_v/F_m) during summer: horizontal leaves were highly photoinhibited whereas vertical leaf orientation protected the leaves from severe photoinhibition. The importance of leaf orientation within the canopy was analysed in two Mediterranean macchia species with distinct strategies for drought and photoinhibition avoidance during summer. The semi-deciduous species (Cistus monspeliensis) exhibited strong seasonal but minimal spatial variability in leaf orientation. Reversible structural regulation of light interception by vertical leaf orientation during summer protected the leaves from severe photoinhibition. The evergreen sclerophyll (Quercus coccifera) exhibited high spatial variability in leaf angle distribution throughout the year and was less susceptible to photoinhibition. The importance of both strategies for plant primary production was analysed with a three-dimensional canopy photoinhibition model (CANO-PI). Simulations indicated that high variability in leaf angle orientation in Q. coccifera resulted in whole-plant carbon gain during the summer, which was 94 % of the maximum rate achieved by theoretical homogeneous leaf orientations. The high spatial variability in leaf angle orientation may be an effective compromise between efficient light harvesting and avoidance of excessive radiation in evergreen plants and may optimize annual primary production. Whole plant photosynthesis was strongly reduced by water stress and photoinhibition in C. monspeliensis; however, the simulations indicated that growth-related structural regulation of light interception served as an important protection against photoinhibitory reduction in whole-plant carbon gain.     

Effect of hydrogen peroxide on the uptake of chlordane by Cucurbita pepo

Background and aims

Plant species can have a major effect on erosion dynamics and soil losses by retaining sediment transported during concentrated runoff. Identifying plant functional traits that influence and predict a species ability for sediment trapping is therefore of great interest, especially to improve management and restoration of degraded lands.

Methods

Sediment trapping ability of four morphologically contrasted species, the broadleaf species Buxus sempervirens and Lavandula angustifolia, and the coniferous species Juniperus communis and Pinus nigra, were investigated with flume experiments. Six functional traits describing stem, leaf and the overall plant morphology, were measured on seedlings. Analyses were performed to compare species efficiency in sediment trapping and to identify traits related to the amount of sediment trapped.

Results

Sediment trapping (RTS) was the highest upslope of Lavandula and the lowest upslope of Juniperus. Principal component analysis showed that RTS was best correlated (positively) with canopy density, described by plant biomass and leaf area per unit volume of plant. Leaf area and plant roundness were also positively related to RTS but to a lesser extent.

Conclusions

The results of this experimental study suggest that canopy completeness, leaf morphology and plant shape influence sediment retention by plants. Such knowledge may improve the diagnosis of land vulnerability to erosion and the prediction of ecosystem functioning after ecological restoration by the construction of bioengineering works in gully floors.     

Two different strategies of Mediterranean macchia plants to avoid photoinhibitory damage by excessive radiation levels during summer drought

The adaptive strategies to high radiation and water stress of the drought tolerant evergreen sclerophylls Quercus coccifera and Arbutus unedo are compared to those of the semi-deciduous Cistus spp. (C. albidus and C. monspeliensis). Cistus spp. partially avoided drought by a marked reduction of their transpirational surface through leaf abscission during summer, when predawn water potential declined below -5.5 MPa. Chlorophyll fluorescence measurements revealed a reversible diurnal decrease of maximum photochemical efficiency of PSII (F_v/F_m), which became more accentuated during summer drought in all species. An important strategy to avoid damage by excessive radiation levels in Cistus spp. was the structural regulation of light interception through leaf angle changes, from a more horizontal orientation in spring (< 35°) to a more vertical orientation in summer (> 70°). Horizontal orientated leaves were highly susceptible to photoinhibition, and excessive radiation often resulted in irreversible photodamage followed by leaf abscission during summer, whereas vertical leaf orientation appeared to protect the leaf from severe photoinhibition. Still, these mechanisms were not fully successful in avoiding chronic photoinhibition, and predawn F_v/F_m values remained low in Cistus spp. during summer (only exhibiting a partial overnight recovery). Evergreen sclerophylls were less susceptible to photoinhibition, and the diurnal decline in F_v/F_m remained fully reversible during drought. Structural regulation of light interception was not found to be an important strategy in these species, and only small, though significant changes in leaf angle occurred. The ecological importance of the adaptive strategies of each functional group is discussed.     

Physiological,morphological and anatomical trait variations between winter and summer leaves of Cistus species

Morphological, anatomical and physiological summer and winter leaf traits of Cistus incanus subsp. incanus, C. salvifolius and C. monspeliensis growing at the Botanical garden of Rome were analyzed. With regard to differences between summer and winter leaves of the considered species, leaf thickness (L) was 21% higher in summer than in winter leaves (mean of the considered species) and this increase was mostly the result of the increased palisade parenchyma thickness over the spongy parenchyma one (24 and 16% higher in summer than in winter leaves, respectively). Leaf mass area (LMA) and leaf tissue density (LTD) were 38% and 17% higher in summer than in winter leaves, respectively (mean of the considered species). The photosynthetic rate (P_N), stomatal conductance (g_s) and chlorophyll content (Chl) of summer leaves were 54%, 17% and 14% lower, respectively, than in winter leaves. C. monspeliensis summer leaves had the highest LMA, LTD, adaxial cuticle thickness (14.6 ± 1.8 mg cm⁻², 1091 ± 94 mg cm⁻³, and 5.8 ± 1.7 μm, respectively) and the lowest mesophyll intercellular spaces (f_ias 38 ± 3%). Moreover, C. monspeliensis had the highest P_N in summer (2.6 ± 0.1 μmol m⁻² s⁻¹) and C. incanus the highest P_N and WUE (84% and 59% higher than the other species) in the favorable period, associated to a higher f_ias (42 ± 2%). C. salvifolius had the highest P_N (54% higher than the other species) in winter. The plasticity index could allow a better interpretation of the habitat preference of the considered species. The physiological plasticity (PI_p = 0.39, mean value of the considered species) was higher than the morphological (PI_m = 0.22, mean value) and anatomical (PI_a = 0.13, mean value) plasticity. Moreover, among the considered species, C. salvifolius and C. incanus are characterized by a larger PI_a (0.14, mean value) which seems to be correlated with their wider ecological distribution and the more favorable conditions of the environments where they naturally occur. The highest PI_m (0.29) of C. monspeliensis indicates that it can play a high adaptive role in highly stressed environments, like fire degraded Mediterranean areas in which it occurs.     

Seasonal Differences in Photochemical Efficiency and Chlorophyll and Carotenoid Contents in Six Mediterranean Shrub Species under Field Conditions

The effects of summer and winter stress on the chlorophyll and carotenoid contents and photosystem 2 efficiency were examined in six Mediterranean scrub species. These six species belong to two different plant functional types: drought semi-deciduous (Halimium halimifolium L., Rosmarinus officinalis L., Erica scoparia L.) and evergreen sclerophylls (Juniperus phoenicea L., Pistacia lentiscus L., Myrtus communis L.). Two sites with different water availability were chosen. In the xerophytic site, despite they belong to two different functional types, R. officinalis and J. phoenicea showed a similar response. These were the most affected species in summer. H. halimifolium showed optimal values of F_v/F_m and non-significant seasonal changes in xanthophyll content. In the mesic site, E. scoparia and M. communis were apparently the most affected species by winter climatic conditions. P. lentiscus presented a pattern similar to H. halimifolium, except for elevated F₀ values. In all the studied species, lutein plus zeaxanthin content was negatively correlated with F_v/F_m in summer and with leaf water potential, thus indicating that the thermal dissipation of energy was a general pattern for all species. Under stress, plant response is more species-specific than dependent on its functional type.     

Drought responses,phenotypic plasticity and survival of Mediterranean species in two different microclimatic sites

• Climate models predict a further drying of the Mediterranean summer. One way for plant species to persist during such climate changes is through acclimation. Here, we determine the extent to which trait plasticity in response to drought differs between species and between sites, and address the question whether there is a trade‐off between drought survival and phenotypic plasticity.
• Throughout the summer we measured physiological traits (photosynthesis – A_max, stomatal conductance – g_s, transpiration – E, leaf water potential – ψl) and structural traits (specific leaf area – SLA, leaf density – LD, leaf dry matter content – LDMC, leaf relative water content – LRWC) of leaves of eight woody species in two sites with slightly different microclimate (north‐ versus south‐facing slopes) in southern Spain. Plant recovery and survival was estimated after the summer drought period.
• We found high trait variability between species. In most variables, phenotypic plasticity was lower in the drier site. Phenotypic plasticity of SLA and LDMC correlated negatively with drought survival, which suggests a trade‐off between them. On the other hand, high phenotypic plasticity of SLA and LDMC was positively related to traits associated with rapid recovery and growth after the drought period.
• Although phenotypic plasticity is generally seen as favourable during stress conditions, here it seemed beneficial for favourable conditions. We propose that in environments with fluctuating drought periods there can be a trade‐off between drought survival and growth during favourable conditions. When climate become drier, species with high drought survival but low phenotypic plasticity might be selected for.

     

Variability in water use efficiency at the leaf level among Mediterranean plants with different growth forms

Assessing natural variability of leaf water use efficiency in plants adapted to extreme conditions of the Mediterranean climate represents an important step in the evaluation of the usefulness of some plant ecophysiological traits under water stress. Eleven Mediterranean species naturally inhabiting the Balearic Islands and corresponding to different growth forms (herbs, semi-deciduous shrubs, woody evergreen shrubs and woody evergreen semi-shrubs) were subject to progressive soil water depletion. Leaf intrinsic water use efficiency was measured by gas exchange at four different degrees of water stress. Under well watered conditions, differences in leaf intrinsic water use efficiency (A _N/g _s) among growth forms were limited to woody evergreen semi-shrubs, which presented the highest values. Under water stress conditions, differences became more evident, with a trend for an increase in A _N/g _s from woody evergreen shrubs, through semi-deciduous shrubs and herbaceous to woody evergreen semi-shrubs. The observed variation in A _N/g _s correlated with several physiological (leaf water potential, soil to leaf hydraulic conductance and stomatal conductance) and morphological (stomatal density) parameters, displaying a general relationship for all growth forms. This suggests that the capacity for withstanding water limitation is adaptive for all Mediterranean species. However, when A _N/g _s was related to leaf mass area, this relationship was not generally applicable, and depended on growth forms, suggesting that different growth forms display specific morphological adjustments in response to water shortage.     

Wood anatomy and hydraulic architecture of stems and twigs of some Mediterranean trees and shrubs along a mesic-xeric gradient

Woody species populating the major Mediterranean ecosystems in the world are characterized by different levels of adaptation to the seasonal Mediterranean climate conditions. Many species of these ecosystems show wood features that allow high efficiency of transport when water is available, while maintaining hydraulic safety during drought periods. This study focuses on the anatomy of juvenile and mature wood of some species representative of continuous sequences of Mediterranean vegetation formations according to gradients of water availability, from xeric to relatively mesic: Cistus monspeliensis L., Rhamnus alaternus L., Myrtus communis L., Pistacia lentiscus L., Olea europaea L., Quercus ilex L., Fraxinus ornus L. and Ostrya carpinifolia L. Twigwood collected in Southern Italy was anatomically compared with the stemwood of the same species represented in the reference slide collection of the National Herbarium of the Netherlands (Lw). The “hydraulic distance” between the wood of main stems and twigs was estimated on the basis of suites of anatomical features related to water efficiency/safety. Although some attributes (i.e. porosity and type of imperforate tracheary elements) were similar in young twigs and older rings, other traits (i.e. vessel frequency and size) evidenced the different hydraulic properties of twig and stemwood. The difference between juvenile and mature structures was large in the species of the mesic end of the gradient while it was relatively small in those more xeric. This tendency is in agreement with the habit gradient from medium-sized trees to small evergreen/drought deciduous shrubs according to decreasing water availability in Mediterranean vegetation types.     

Leaf litter of <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) inhibits seed germination in sclerophyllous montane woodlands of central Chile

Leaf litter accumulation can have either positive, negative or neutral effects on seed germination and seedling recruitment. In montane woodlands of the Mediterranean zone of central Chile, large amounts of leaf litter accumulate beneath the crowns of the summer semi-deciduous tree Kageneckia angustifolia and no regeneration of this or other plant species has been observed beneath this tree throughout the year. In a sample plot of 5000 m² we selected ten K. angustifolia trees and measured (1) leaf litter accumulation beneath and outside canopy; (2) the effects of time elapsed since burial on viability of K. angustifolia seeds with and without a leaf litter cover; (3) field seed germination with presence or absence of leaf litter and (4) the possible chemical effects of K. angustifolia leaf litter leachates on seed germination of its own seeds and of other two co-occurring native shrubs species (Guindilia trinervis and Solanum ligustrinum). Our results show that a considerable accumulation of leaf litter occurred beneath K. angustifolia, and litter negatively affected seed viability and germination of this species in the field. Under laboratory conditions, K. angustifolia leaf litter leachates inhibited seed germination of its own seeds and of the two native shrub species. Chemical effects are likely involved in the negative effects of leaf litter on the recruitment of K. angustifolia in the montane sclerophyllous woodland of central Chile.     

Sex-related differences in leaf traits in an androdioecious shrub under contrasting levels of soil salinity

This study explores whether male and hermaphrodite plants of Phillyrea angustifolia (Oleaceae) show physiological and structural differences at the leaf level under severe water stress driven by drought and soil salinity. Leaf traits were measured in summer, at the height of the summer drought period, in male and hermaphrodite plants from two adjacent sites under contrasting soil salinity levels. Male plants from the saline site had significantly higher leaf proline content compared to males from the nonsaline site. By contrast, leaf proline levels were similarly low in hermaphrodite plants from both sites. On the other hand, hermaphrodite plants from the saline site had higher leaf stomatal frequency than hermaphrodites from the nonsaline site, whereas this parameter did not differ for male plants across sites. Such differences could be interpreted as the result of two different solutions to the same selective pressure in the androdioecious shrub P. angustifolia.     

Leaf traits and relationships differ with season as well as among species groupings in a managed Southeastern China forest landscape

The relationships among leaf traits often reflect plant adaptation for coping with nutrient resources. However, the seasonal variations in leaf traits and their relationship with soil nutrients are not well understood. We sampled seven major functional traits of thirty trees and nine shrubs (sorted into different plant functional groups, PFGs, based on their growth form, leaf lifespan, and leaf shape) at different seasons in a managed forest plantation of Southeastern China. Both green leaf nitrogen and phosphorus concentrations (N_green and P_green) decreased significantly from spring and summer to autumn, and varied significantly with PFGs (P?<?0.05) at different times of the year. Across all plants, specific leaf area correlated positively with N_green and P_green in spring, summer, and winter, but not in autumn; N resorption proficiency generally correlated positively with N_green in each season, while P resorption efficiency correlated positively with P_green in spring and summer, but not in autumn and winter. Soil nitrogen availability correlated negatively with leaf nutrient traits in some seasons. In conclusion, leaf trait relationships varied among the seasons and among PFGs. Seasonal dynamics of leaf traits as well as soil nutrients?? relations must be considered when exploring plant feedback to soil nutrients.     

Growth, water relations and photosynthesis of seedlings and resprouts after fire

Seasonal patterns of growth, water relations, photosynthesis and leaf characteristics were compared between obligate seeders (Cistus monspeliensis and Cistus ladanifer) and resprouters (Arbutus unedo and Pistacia lentiscus) from the first to the second year after fire. We hypothesized that seedlings would be more water-limited than resprouts due to their shallower root systems. Regarding water use strategies, Cistus species are drought semi-deciduous and A. unedo and P. lentiscus are evergreen sclerophylls, therefore, comparisons were based on the relative deviation from mature conspecific plants. Seedlings and resprouts had higher shoot elongation and leaf production than mature plants, and over an extended period. Differences from mature plants were larger in resprouts, with two-fold transpiration, leaf conductance and photosynthesis in late spring/early summer. Seedlings of C. monspeliensis exhibited higher transpiration and leaf conductance than mature plants, while those of C. ladanifer only exhibited higher water potential. Growth increments and ameliorated water relations and photosynthesis after fire were attributed to an increase in water and nutrient availability. The small differences in water relations and photosynthesis between seedlings and mature conspecifics are in accordance with the prediction of seedlings experiencing higher water limitation than resprouts. We attribute these results to differences in root systems: resprouters benefited from an increase in root/shoot ratios and the presence of deep roots whereas Cistus seedlings relied on very shallow roots, which cannot provide assess to deep water during summer. Nevertheless, seedlings did not show evidence of experiencing a more severe water limitation than mature conspecifics, which we attributed to the presence of efficient mechanisms of avoiding and tolerating water stress. The results are discussed in relation to post-fire demography of seeders and resprouters in Mediterranean communities.     

Summer-drought constrains the phenology and growth of two coexisting Mediterranean oaks with contrasting leaf habit: implications for their persistence and reproduction

This study analyses how coexisting evergreen and deciduous oaks adjust their phenology to cope with the stressful Mediterranean summer conditions. We test the hypothesis that the vegetative and reproductive growth of the winter deciduous (Quercus faginea Lam.) is more affected by summer drought than that of the evergreen [Quercus ilex L. subsp. ballota (Desf.) Samp.]. First, we assessed the complete aboveground phenology of both species during two consecutive years. Shoot and litter production and bud, acorn and secondary growth were monitored monthly. Second, we identified several parameters affected by summer conditions: apical bud size, individual leaf area (LA), leaf mass per area (LMA) and acorn yield in both species, and leaf-fall in Q. faginea; and analysed their variation over 10 years. Q. ilex performed up to 25% of shoot growth and most leaf development during summer, whereas Q. faginea completed most of both phenophases during spring. Secondary growth was arrested in summer under drought conditions. Approximately, 30–40% of bud and 40–50% of acorn growth was undertaken during summer in both species. Summer drought related to differences in LA, LMA and leaf senescence, but not to acorn yield. Both species had similar year-to-year patterns of acorn production, though yields were always lower in Q. faginea. Bud size decreased severely in both species during extremely dry years. In Q. ilex, bud size tended to alternate between years of large and small buds, and these patterns were followed by opposite trends in stem length. In Q. faginea, bud size was more stable through time. Q. ilex was more phenologically active during summer than Q. faginea, indicating a higher tolerance to drought. Furthermore, bud and fruit growth (the only two phenophases that both species performed during summer) were more severely affected by summer drought in Q. faginea than in the evergreen. The differential effects of summer drought on key phenophases for the persistence (bud growth) and colonization ability (fruit production) of both species may have consequences for their coexistence.     

Seasonality and phenology alter functional leaf traits

In plant ecophysiology, functional leaf traits are generally not assessed in relation to phenological phase of the canopy. Leaf traits measured in deciduous perennial species are known to vary between spring and summer seasons, but there is a knowledge gap relating to the late-summer phase marked by growth cessation and bud set occurring well before fall leaf senescence. The effects of phenology on canopy physiology were tested using a common garden of over 2,000 black cottonwood (Populus trichocarpa) individuals originating from a wide geographical range (44–60ºN). Annual phenological events and 12 leaf-based functional trait measurements were collected spanning the entire summer season prior to, and following, bud set. Patterns of seasonal trait change emerged by synchronizing trees using their date of bud set. In particular, photosynthetic, mass, and N-based traits increased substantially following bud set. Most traits were significantly different between pre-bud set and post-bud set phase trees, with many traits showing at least 25 % alteration in mean value. Post-bud set, both the significance and direction of trait–trait relationships could be modified, with many relating directly to changes in leaf mass. In Populus, these dynamics in leaf traits throughout the summer season reflected a shift in whole plant physiology, but occurred long before the onset of leaf senescence. The marked shifts in measured trait values following bud set underscores the necessity to include phenology in trait-based ecological studies or large-scale phenotyping efforts, both at the local level and larger geographical scale.     

Differential impact of liana colonization on the leaf functional traits of co-occurring deciduous and evergreen trees in a tropical dry scrub forest

The present study was carried out to analyze the leaf functional traits of co-occurring evergreen and deciduous tree species in a tropical dry scrub forest. This study also intended to check whether the species with contrasting leaf habits differ in their leaf trait plasticity, responding to the canopy infestation by lianas. A total of 11 leaf functional traits were studied for eight tree species with contrasting leaf habits (evergreen and deciduous) and liana-colonization status (with or without liana). In the liana-free environment (L^–), evergreen trees had significantly higher leaf tissue density (LTD) and total chlorophyll (CHL_t) than the deciduous species. Whereas the deciduous trees had higher specific leaf area (SLA) and mass-based leaf nitrogen concentration (N_mass). The leaf trait-pair relationship in the present study agreed with the well-established global trait-pair relationships (leaf thickness (LT) vs. SLA, N_mass vs. LT, SLA vs. N_mass, and LDMC vs. SLA). There was a significant difference between L⁺ and L^– individuals in leaf area (LA), petiole length (PL), SLA, LDMC, and CHL_t in the deciduous species. On the other hand, evergreen species showed marked differences across LT, SLA, LTD, N_mass, and chlorophyll components between L⁺ and L^– individuals of the same species. The results revealed the differential impact of liana colonization on the host trees with contrasting leaf habits. The deciduous species with the acquisitive strategy can have a competitive advantage over evergreen species in the exposed environments (L^–), whereas evergreen species with shade-tolerant properties were better acclimated to the shaded environments (L⁺). Therefore, liana colonization can significantly impact the C-fixation strategies of the host trees by altering their light environment and further, the magnitude of such impact may vary among species of different leaf habits. The result also indicated the patterns of convergence and divergence in some of the leaf functional traits between evergreen and deciduous species explaining the patterns of species co-existence.

     

Earlier summer drought affects leaf functioning of the Mediterranean species Cistus monspeliensis L.

The Mediterranean vegetation is characterized by a high diversity of growth forms, habits and phenology that enable it to endure under harsh environmental conditions. It is however unclear whether these adaptations may allow plant survival under more extreme conditions, as predicted by climatic models under the perspective of climate change. A manipulative experiment aiming at anticipating summer aridity has been run to analyse the effects of the experimental drought on spring-leaf functioning and characteristics of the leaf-dimorphic Mediterranean shrub Cistus monspeliensis L.Assimilation rates were reduced under anticipated summer aridity due to a decrease of stomatal conductance, but only before morphological adaptations to drought (an increase of leaf mass per area) occurred. These adaptations were anticipated under experimental dry conditions, and causes photosynthetic performances to recover compared to previous dates. When natural summer aridity occurred, the leaf mass per area also changed in the control. However, this causes no recovery of the photosynthetic performances, because of the decrease of stomatal conductance due to low soil water content and leaf water potential values. Moreover, under experimental drought, leaf shedding was anticipated to reduce water losses, causing an overall reduction of leaf lifespan.     

Seasonal physiological plasticity and recovery capacity after summer stress in Mediterranean scrub communities

Under natural conditions the overlapping of multiple stressors may initiate coordinated ecophysiological responses in Mediterranean species. Seasonal plasticity may enable plants to better cope with adverse environmental conditions and/or resource variability. In this article, we study the seasonal responses of 12 woody species in two sites of differing water availability, in a Mediterranean-type climate. Plants were measured for water potential, photochemical efficiency, photosynthetic pigments and leaf proline content throughout the year. The results revealed that species presented different ecophysiological strategies, even when sharing the same area. In the xerophytic site, some species suffered severe water stress (−12 MPa and F _v/F _m lower than 0.3), while others exhibited optimal values of F _v/F _m and only moderate water stress. All the plants recovered after the first autumn rains. In the hygrophytic site, some sclerophyll species did not exhibit signs of water stress, but did suffer photoinhibition in clear winter days. A plasticity index was calculated to provide an integrated value of species plasticity. In summer, plasticity was higher in the xerophytic site, while in winter it was higher in the hygrophytic site. Ordination analysis of the physiological traits supports the traditional gradient of Mediterranean strategies from drought semideciduous to evergreen sclerophyll species, although spiny legume species formed an independent functional group. The functional responses of species clearly differ among plant communities according to the prevailing site stressors, but no unique pattern emerges. Species combine traits in broader strategies according to previous evolutionary story exhibiting a certain amount of trade among traits, each contributing to alleviate a part of the plant stress.     

柯-青冈常绿阔叶林优势树种叶片性状变异及适应策略

厘清叶片功能性状的变异及性状间的权衡关系,对揭示植物对环境变化的响应及适应策略具有重要意义。以中亚热带柯(Lithocarpus glaber)-青冈(Cyclobalanopsis glauca)常绿阔叶林为对象,测量了1 hm²固定监测样地内6个优势树种的叶面积(LA)、比叶面积(SLA)、干物质含量(LDMC)、叶片厚度(LT)、叶片碳(LC)、叶片氮(LN)、叶片磷(LP)含量和叶片碳氮比(LC ∶ LN)8个功能性状,采用多重比较、主成分分析(PCA)法分析了群落叶片功能性状的生活型、种内和种间变异及性状间关系。结果表明：(1)群落内叶片功能性状种内、种间差异显著,变异中等(CV： 0.02-0.59),其结构性状的可塑性较化学性状保守,变异格局符合"性状空间变异分割假说";针叶树种的LA、SLA显著低于阔叶树种,常绿树种的LC和LDMC最小,而落叶树种的SLA、LN和LP最大以及LT和LC ∶ LN最小。(2)群落叶片功能性状变异主要来源于生活型和种间变异,种内变异亦有显著贡献;生活型对多数性状的贡献率较大,其对LDMC、LC ∶ LN的贡献分别高达93.11%和91.76%;种间变异(LDMC除外)对结构性状的贡献率均高于化学性状;种内变异对LP的贡献率(23.66%)较种间变异高。(3)叶片性状之间多表现出显著相关关系,针叶树和阔叶树在PCA排序图中聚集于不同区域,叶经济型谱在柯-青冈群落中具有适用性。不同树种通过叶片结构、化学性状之间的权衡策略来适应环境变化,从而实现群落物种共存。结果可为理解森林群落物种的环境适应策略、预测群落动态变化和制定植被恢复措施提供科学依据。     

Foliage dynamics, leaf traits, and growth of coexisting evergreen and deciduous trees in a tropical montane forest in Ethiopia

Foliage dynamics of three functional tree types representing major components of the tropical montane evergreen forest in southern part of Central Ethiopia were compared. The species were Podocarpus falcatus (evergreen gymnosperm), Prunus africana (evergreen broadleaf), and Croton macrostachyus (facultative deciduous). The hypothesis examined is that in such tropical trees, endogenous control of foliage dynamics by the leaf life-spans (LLS) is largely dominant over external signals. Crown foliage turnover, leafiness of twigs, LLS, photosynthetic performance, respiration rate, specific leaf area, and relative growth rates of the stems were investigated. Foliage dynamics and leafiness of the twigs were monitored over 2?years while leaf traits were followed over 3?months. The degree of inter and intra-individual synchronization of foliage phenophases was examined to get an estimate of the contributions of endogenous and external signals to the dynamics of the foliages. Autoregression analysis indicated significant influence of the moisture regime on leaf sprouting of Croton and Podocarpus. During pronounced dry periods, new leaves were not developed. Analysis of phenological data using circular statistics revealed that in spite of strong inter-individual synchronization of leaf flush and fall (Podocarpus and Croton), the dynamics of individual parts of the crowns were less synchronized. LLS was independent of climate factors and it had substantial contribution to the control of foliage turnover. Moreover, examination of ecophysiological traits of developing leaves of the studied functional types showed differing patterns with LLS corroborating the ecophysiological characteristics. Although overlaid by fungal infestation, both the foliage and ecophysiological properties of Prunus resemble that of Podocarpus but the former exhibited a shorter LLS and slightly higher metabolic rates. Nevertheless, all species reacted positively to high moisture with respect to stem growth. In spite of largely differing weather conditions of the 2?years, direct competitive advantage of one of the species over the others could not be detected.     

How do functional traits syndromes covary with growth and reproductive performance in a water‐stressed population of Fagus sylvatica?

A central issue in plant evolutionary ecology is to understand how several coordinated suites of traits (i.e. traits syndrome) may be jointly selected within a single species. This study aims to describe patterns of variation and co‐variation of functional traits in a water‐stressed tree population and test their relationships with performance traits. Within a Mediterranean population of Fagus sylvatica experiencing recurrent summer droughts, we investigated the phenotypic variation of leaf unfolding phenology, leaf area (LA), leaf mass per area (LMA), leaf water content (LWC), water use efficiency (WUE) estimated by carbon isotopic discrimination (d13C), twig Huber‐value (HV: the stem cross‐section divided by the leaf area distal to the stem), wood density (WDens), and leaf nitrogen content (N_mass). First, a principal component analysis revealed that two main axes structured the phenotypic variability: the first axis opposed leaf unfolding earliness and LWC to LMA and WUE; the second axis opposed LA to HV. These two axes can be interpreted as the opposition of two strategies (water economy versus water uptake) at two distinct scales (leaf for the first axis and branches for the second axis). Second, we found that LMA, LA, leaf unfolding and LWC responded differently to competition intensity, while WUE, WDens and HV did not correlate with competition. Third, we found that all studied functional traits were related to growth and/or reproductive performance traits and that these relationships were frequently non‐linear, showing strong interactions between traits. By highlighting phenotypic clustering of functional traits involved in response to water stress and by evidencing antagonistic selection favouring intermediate trait values as well as trait combinations, our study brought new insights on how natural selection operates on plant functional traits in a stressful environment.