Relationship between net photosynthesis and leaf respiration in Mediterranean evergreen species

The relationship between net photosynthetic (P _N) and leaf respiration (R) rates of Quercus ilex, Phillyrea latifolia, Myrtus communis, Arbutus unedo, and Cistus incanus was monitored in the period February 2006 to February 2007. The species investigated had low R and P _N during winter, increasing from March to May, when mean air temperature reached 19.2 °C. During the favourable period, C. incanus and A. unedo had a higher mean P _N (16.4±2.4 μmol m⁻² s⁻¹) than P. latifolia, Q. ilex, and M. communis (10.0±1.3 μmol m⁻² s⁻¹). The highest R (1.89±0.30 μmol m⁻² s⁻¹, mean of the species), associated to a significant P _N decrease (62 % of the maximum, mean value of the species), was measured in July (mean R/P _N ratio 0.447±0.091). Q₁₀, indicating the respiration sensitivity to short-term temperature increase, was in the range 1.49 to 2.21. Global change might modify R/P _N determining differences in dry matter accumulation among the species, and Q. ilex and P. latifolia might be the most favoured species by their ability to maintain sufficiently higher P _N and lower R during stress periods.     

Morphological and chemical leaf composition of Mediterranean evergreen tree species according to leaf age

Changes in morphology [leaf dry mass per unit area (LMA), thickness and density] and chemical composition (macronutrients and fibres content) in different age leaves of eight evergreen Mediterranean woody species were investigated. LMA and leaf thickness increased with leaf age increasing. Young tissues possessed higher concentrations of N, P, K, and Mg and lower Ca concentrations on a dry mass basis. However, mineral content was independent of age on leaf area basis (except for Ca content) suggesting that the changes in mineral concentration with leaf ageing are due to dilution in the larger dry mass accumulated in the oldest leaves. Leaf tissue density (LTD) increased during the first year of the leaf life. Lignin and hemicellulose concentrations did not vary along leaf life and the cellulose concentration increased with leaf age in most species between the current-year and the one-year old leaves. Our results suggested that physical leaf reinforcement with a higher cellulose concentration and density might be a leaf response to the unfavourable climatic conditions during the first winter.     

Long-time variations in leaf mass and area of Mediterranean evergreen broad-leaf and narrow-leaf maquis species

Morphological (dry mass, DM; surface area, LA; leaf mass per area, LMA), anatomical (leaf thickness, L), phenological (leaf life span, LL), and physiological (net photosynthetic rate, P _N) leaf traits of the evergreen species co-occurring in the Mediterranean maquis developing at Castelporziano (Rome) were tested. The correlation analysis indicated that LMA variation was tightly associated with LL variations: Cistus incanus and Arbutus unedo had a short LL (4±1, summer leaves, and 11±1 months, respectively) and low LMA (153±19 g m⁻²) values, Quercus ilex, Phillyrea latifolia, and Pistacia lentiscus high LMA (204±7 g m⁻²) and long LL (22±3 months), Erica arborea, Erica multiflora, and Rosmarinus officinalis a short LL (9±2 months) and an either high (213±29 g m⁻², R. officinalis and E. multiflora) or low (115±17 g m⁻², E. arborea) LMA. LMA values were significantly (p≤0.05) correlated with P _N (r≥0.68). In the tested species, LMA increased in response to the decrease of the total rainfall during the leaf expansion period. LMA variation was due to the unequal variation of DM and LA in the considered species. LMA is thus a good indicator of evergreen maquis species capability to respond to climate change, in particular to total rainfall decrease in the Mediterranean basin.     

Allozyme characterisation of four Mediterranean evergreen oak species

Polymorphism at 11 enzyme loci was used to compare the four evergreen oak species of the Mediterranean Basin and to establish their taxonomical relationships. As a comparison, two evergreen oak species from Tibet, located in the primary distribution area of evergreen oaks, were analysed at the same loci. Cladistic analyses led to the separation of a cluster of four species, namely the three Mediterranean Q. ilex, Q. coccifera and Q. alnifolia and the Tibetan Q. aquifolioides. The other Tibetan species, Q. semicarpifolia, and Q. suber, from the Western Mediterranean Basin, were very distinct genetically from the four other species. These results were not consistent with previous taxonomical treatment, based on morphology alone, which classified Q. ilex and Q. coccifera in “ilex group” (=subgenus Sclerophyllodrys O. Schwartz), and the other four species in “cerris group” (subgenus Cerris, according to Schwartz). Allozyme variation has thus provided useful information to clarify the taxonomy of evergreen oaks.     

Allometric models for leaf area estimation across different leaf-age groups of evergreen broadleaved trees in a subtropical forest

The accurate and nondestructive determination of individual leaf area (LA) of plants, by using leaf length (L) and width (W) measurement or combinations of them, is important for many experimental comparisons. Here, we propose reliable and simple regressions for estimating LA across different leaf-age groups of eight common evergreen broadleaved trees in a subtropical forest in Gutianshan Natural Reserve, eastern China. During July 2007, the L, W, and LA of 2,923 leaves (202 to 476 leaves for each species) were measured for model construction and the respective measurements on 1,299 leaves were used for model validation. Mean L, W, LA and leaf shape (L:W ratio) differed significantly between current and older leaves in four out of the eight species. The coefficients of one-dimension LA models were affected by leaf age for most species while those incorporating both leaf dimensions (L and W) were independent of leaf age for all the species. Therefore, the regressions encompassing both L and W (LA = a L W + b), which were independent of leaf age and also allowed reliable LA estimations, were developed. Comparison between observed and predicted LA using these equations in another dataset, conducted for model validation, exhibited a high degree of correlation (R ² = 0.96−0.99). Accordingly, these models can accurately estimate the LA of different age groups for the eight evergreen tree species without using instruments.     

Divergent scaling of respiration rates to nitrogen and phosphorus across four woody seedlings between different growing seasons

Empirical studies indicate that the exponents governing the scaling of plant respiration rates (R) with respect to biomass (M) numerically vary between three‐fourth for adult plants and 1.0 for seedlings and saplings and are affected by nitrogen (N) and phosphorus (P) content. However, whether the scaling of R with respect to M (or N and P) varies among different phylogenetic groups (e.g., gymnosperms vs. angiosperms) or during the growing and dormant seasons remains unclear. We measured the whole‐plant R and M, and N and P content of the seedlings of four woody species during the growing season (early October) and the dormant season (January). The data show that (i) the scaling exponents of R versus M, R versus N, and R versus P differed significantly among the four species, but (ii), not between the growing and dormant seasons for each of the four species, although (iii) the normalization constants governing the scaling relationships were numerically greater for the growing season compared to the dormant season. In addition, (iv) the scaling exponents of R versus M, R versus N, and R versus P were numerically larger for the two angiosperm species compared to those of the two gymnosperm species, (v) the interspecific scaling exponents for the four species were greater during the growing season than in the dormant season, and (vi), interspecifically, P scaled nearly isometric with N content. Those findings indicate that the metabolic scaling relationships among R, M, N, and P manifest seasonal variation and differ between angiosperm and gymnosperm species, that is, there is no single, canonical scaling exponent for the seedlings of woody species.     

降雨量改变对常绿阔叶林干旱和湿润季节土壤呼吸的影响

通过野外原位试验,研究降雨量改变对华西雨屏区常绿阔叶林干旱和湿润季节土壤呼吸速率的影响。采用LI-8100土壤碳通量分析系统(LI-COR Inc.,USA)测定干旱和湿润季节对照(CK)、增雨10%(LA)、增雨5%(TA)、减雨10%(LR)、减雨20%(MR)、减雨50%(HR)6个处理水平的土壤呼吸速率,并通过回归方程分析温度和湿度与土壤呼吸速率间的关系。结果表明:湿润季节土壤呼吸速率高于干旱季节,HR处理对干旱季节土壤呼吸速率影响较大,而LA处理对湿润季节土壤呼吸速率的影响较大。TA和LR处理使土壤呼吸的温度敏感性增加,而HR、LA和MR处理使土壤呼吸的温度敏感性降低,干旱季节Q10值高于湿润季节。各处理湿润季节土壤微生物量碳氮含量显著高于干旱季节,HR、MR和LA处理减少土壤微生物生物量碳、氮的含量,而TA和LR处理增加土壤微生物生物量碳、氮的含量。与湿润季节相比,干旱季节土壤水分对土壤呼吸速率的影响较大;而与土壤温度相比,土壤水分对土壤呼吸速率的影响较小。在降雨量改变的背景下,华西雨屏区常绿阔叶林无论是干旱还是湿润季节,适当增雨和减雨都会促进土壤呼吸速率,而较高量的增雨和减雨会抑制土壤呼吸速率。     

Root respiration and associated costs in deciduous and evergreen species of Quercus

Root construction and maintenance costs were estimated in four evergreen and three deciduous Quercus species that are typical in the landscape of southern Spain. The cost quantification was based on analysis of the growth–­respiration ratio. Values observed for both construction cost (ranging from 1·17 to 1·29 g glucose g^?1 dry weight) and maintenance cost (ranging from 6·22 to 11·71 mg glucose g^?1 dry weight d^?1) were generally lower than those reported in other studies. The results showed non‐significant differences between deciduous and evergreen species. The lack of significant differences between species appeared to be due to the homogeneity of growth conditions. Hydroponic culture, with unrestricted nutrient and water supply, would lead to low tissue carbon content and low respiration rates, leading to the low costs observed. Furthermore, the fact that root organs are clearly importers of organic molecules inevitably entails some underestimation of the respiration associated with growth and, to a lesser extent, with maintenance respiration. This leads in turn to underestimation of the corresponding construction and maintenance costs. All this raises doubts as to the suitability of this method for studying root systems.     

亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系,其异速生长指数<1.0,但该假说是否适用于不同生活型(常绿与落叶)亚热带木本植物不同冠层高度(上下冠层)当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象,采用标准化主轴回归估计(standardized major axis estimation, SMA)方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示:(1)当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异(P>0.05),叶面积在常绿和落叶植物中有显著差异(P<0.05),常绿和落叶植物的比叶重存在显著差异(P<0.05),而落叶植物的比叶重在不同冠层高度存在显著差异(P<0.05),同一冠层,常绿植物比叶重显著高于落叶植物(P<0.05);(2)69种植物的叶面积与叶生物量异速生长指数具有物种特异性,60.9%的物种叶面积与叶生物量呈等速生长关系;(3)不同冠层和生活型植物的叶面积与...     

Variation in evergreen and deciduous species leaf phenology in Assam, India

In the present study phenological activities such as leaf and shoot growth, leaf pool size and leaf fall were observed for 3 years (March 2007–March 2010) in 19 tree species (13 evergreen and 6 deciduous species) in a wet tropical forest in Assam, India. The study area receives total annual average rainfall of 2,318 mm of which most rain fall (>70 %) occurs during June–September. Both the plant groups varied significantly on most of the shoot and leaf phenology parameters. In general, growth in deciduous species initiated before the evergreen species and showed a rapid shoot growth, leaf recruitment and leaf expansion compared to evergreen species. Leaf recruitment period was significantly different between evergreen (4.2 months) and deciduous species (6.8 months). Shoot elongation rate was also significantly different for evergreen and deciduous species (0.09 vs. 0.14 cm day^?1 shoot^?1). Leaf number per shoot was greater for deciduous species than for evergreen species (34 vs. 16 leaves). The average leaf life span of evergreen species (328 ± 32 days) was significantly greater than that of deciduous species (205 ± 16 days). The leaf fall in deciduous species was concentrated during the winter season (Nov–Feb), whereas evergreens retained their leaves until the next growing season. Although the climate of the study area supports evergreen forests, the strategies of the deciduous species such as faster leaf recruitment rate, longer leaf recruitment time, faster shoot elongation rate during favorable growing season and short leaf life span perhaps allows them to coexist with evergreen species that have the liberty to photosynthesize round the year. Variations in phenological strategies perhaps help to reduce the competition among evergreen and deciduous species for resources in these forests and enable the coexistence of both the groups.     

Effects of root diameter and root nitrogen concentration on in situ root respiration among different seasons and tree species

Knowledge of root respiration is a prerequisite for a better understanding of ecosystem carbon budget and carbon allocation. However, there are not many relevant data in the literature on direct measurements of in situ root respiration by root chamber method. Furthermore, few studies have been focused on the effects of root diameter (D _r) and root nitrogen concentration (N _r) on in situ root respiration among different seasons and tree species. To address these goals, we used a simplified root-chamber system to measure in situ root respiration rates of Acacia crassicarpa and Eucalyptus urophylla in subtropical plantations of south China. We found that the species and season variation in root respiration were affected by D _r and N _r. Also, the root respiration per unit dry mass (R _r, nmol CO₂ g⁻¹ s⁻¹) and root respiration per unit N (R _n, nmol CO₂ g N⁻¹ s⁻¹) were affected by D _r and N _r. The R _r, R _n, N _r and soil temperature sensitivity (Q ₁₀) of R _r for the two species significantly decreased with an increase of D _r. The R _r of the two species showed significant an inter-seasonal and diurnal pattern, and this trend decreased with increasing D _r. Both the R _r and Q ₁₀ of the two species increased with increasing N _r. The D _r and N _r explained 54 and 52% of the observed variation in R _r for A. crassicarpa, and 65 and 70% for E. urophylla. The R _r, N _r, and Q ₁₀ of A. crassicarpa were significantly higher than those of E. urophylla. Our results indicated that root respiration was dependent on D _r and N _r, and this dependence varied with season and plant species.     

Stomatal patchiness in Mediterranean evergreen sclerophylls

Midday depression of net photosynthesis and transpiration in the Mediterranean sclerophylls Arbutus unedo L. and Quercus suber L. occurs with a depression of mesophyll photosynthetic activity as indicated by calculated carboxylation efficiency (CE) and constant diurnal calculated leaf intercellular partial pressure of CO₂ (C_i). This work examines the hypothesis that this midday depression can be explained by the distribution of patches of either wide-open or closed stomata on the leaf surface, independent of a coupling mechanism between stomata and mesophyll that results in a midday depression of photosynthetic activity of the mesophyll. Pressure infiltration of four liquids differing in their surface tension was used as a method to show the occurrence of stomatal patchiness and to determine the status of stomatal aperture within the patches. Liquids were selected such that the threshold leaf conductance necessary for infiltration through the stomatal pores covered the expected diurnal range of calculated leaf conductance (g) for these species. Infiltration experiments were carried out with leaves of potted plants under simulated Mediterranean summer conditions in a growth chamber. For all four liquids, leaves of both species were found to be fully infiltratable in the morning and in the late afternoon while during the periods leading up to and away from midday the leaves showed a pronounced patchy distribution of infiltratable and non-infiltratable areas. Similar linear relationships between the amount of liquid infiltrated and g (measured by porometry prior to detachment and infiltration) for all liquids clearly revealed the existence of pneumatically isolated patches containing only wide-open or closed stomata. The good correspondence between the midday depression of CE, calculated under the assumption of no stomatal patchiness, and the diurnal changes in non-infiltratable leaf area strongly indicates that the apparent reduction in mesophyll activity results from assuming no stomatal patchiness. It is suggested that simultaneous responses of stomata and mesophyll activity reported for other species may also be attributed to the occurrence of stomatal patchiness. In Quercus coccifera L., where the lack of constant diurnal calculated C_i and major depression of measured CE at noontime indicates different stomatal behavior, non-linear and dissimilar relationships between g and the infiltratable quantities of the four liquids were found. This indicates a wide distribution of stomatal aperture on the leaf surface rather than only wide-open or closed stomata.Dedicated to Professor Otto L. Lange on the occasion of his 65th birthday     

Seasonal variation in leaf characteristics and food selectionby larval noctuids on an evergreen Mediterranean shrub

Despite year round availability of foliage, abundance of generalist noctuid larvae (Lepidoptera: Noctuidae) in evergreen-dominated Mediterranean forests has a narrow, distinct spring peak. This restricted larval period has been suggested to result in part from avoidance of the nutritionally poor mature foliage, and preference for nutritionally superior spring-produced young leaves. This study examines this hypothesis by (i) documenting differences in nutritional characteristics between expanding (April) and mature (June) young leaves of the evergreen Mediterranean shrub Daphne laureola L. (Thymelaeaceae), and (ii) experimentally studying the feeding preferences of noctuid larvae for young leaves, old leaves (≥ 1 yr old), and developing fruits of this species in one south-eastern Spanish locality. Young leaves of D. laureola declined in nutrient concentration and specific dry mass from April to June. The responses of noctuid larvae, in terms of both relative preference and total consumption, to this seasonal variation in chemical and physical features of young leaves were also investigated. When noctuid larvae were simultaneously offered young leaves, old leaves and developing fruits, they exhibited similar preferences for young leaves and developing fruits, and rejected old leaves developed during the previous year. Noctuid larvae did not modify their consumption of young leaves relative to old leaves and developing fruits in response to seasonal changes. Food selection patterns exhibited by D. laureola noctuid herbivores, notably the rejection of old leaves in favour of young ones, are consistent with the hypothesis relating restricted larval periods of these generalist consumers with the low food value of the previous season leaves of evergreen Mediterranean plants.     

Leaf traits and water relations of 12 evergreen species in Costa Rican wet and dry forests: patterns of intra-specific variation across forests and seasons

This study examined variation in leaf traits and water relations in 12 evergreen and semideciduous woody species that occur in both seasonal wet and dry forests in Costa Rica and compared intra-specific leaf–trait correlations to those found in inter-specific global studies. The following traits were measured in both forests across seasons for 2 years: leaf nitrogen (N), leaf carbon (C), specific leaf area (SLA), toughness, cuticle thickness, leaf thickness, and leaf lifespan (LLS). Leaf water potential (LWP) and water content (LWC) were measured as indices of plant available water. Canopy openness, soil moisture, and herbivory were also measured to compare environmental variation across sites. Although species contributed the greatest amount to variation in traits, season, forest, and their interaction had a large influence on patterns of intra-specific leaf–trait variation. Leaf traits that contributed most to variation across sites were C, LWP, leaf thickness, and SLA. Traits that contributed most to variation across seasons were leaf toughness, LWP, and LWC. Furthermore, leaf traits were more correlated (i.e., number and strength of correlations) in the dry than in the wet forest. In contrast to results from global literature syntheses, there was no correlation between LLS and N, or LLS and SLA. Both light and water availability vary seasonally and may be causing variation in a number of leaf traits, specifically those that relate to water relations and leaf economics. Strong seasonality may cause leaf–trait relationships at the local scale to differ from those documented in continental and global-scale studies.     

Linking root respiration to chemistry and morphology across species

Root respiration is a critical physiological trait involved in root resource acquisition strategies, yet it is less represented in root trait syndrome. Here we compiled a large dataset of root respiration associated with root chemical and morphological traits from 245 plant species. Our results demonstrated that root respiration correlated positively with root nitrogen concentration (RNC) and negatively with root tissue density (RTD) across and within woody and non‐woody species. However, the relationships between root respiration and specific root length (SRL) and root diameter (RD) were weak or even insignificant. Such root respiration–traits relationships were not completely in line with predictions by the root economics spectrum (RES). Furthermore, the principal component analysis showed that root trait syndrome was multidimensional. Root respiration was associated more strongly with the RNC‐RTD axis (the classical RES) than with the orthogonal SRL‐RD axis for woody species, but not for non‐woody species. Collectively, the linkages of root physiological, chemical, and morphological traits provide a better understanding of root trait covariation and root resource acquisition strategies.