Renal anatomy in sparrows from different environments

The renal anatomy of three species of sparrows, two from mesic areas, the House Sparrow (Passer domesticus) and Song Sparrow (Melospiza melodia), and one salt marsh species, the Savannah Sparrow (Passerculus sandwichensis) was examined. Electron microscopy was used to describe the ultrastructure of the nephron. In addition, stereology was used to quantify the volumes of cortex, medulla, and major vasculature of the kidneys, and the volumes and surface areas occupied by individual nephron components. There appeared to be no differences in the ultrastructural anatomy of the nephrons among the sparrows. Proximal tubules contained both narrow and wide intercellular spaces filled with interdigitations of the basolateral membrane. The thin limbs of Henle contained very wide intercellular spaces which were absent in the thick limbs of Henle. The distal tubule cells contained short, apical microvilli and infoldings of the basolateral membrane. In cross section, the medullary cones of all birds display an outer ring of thick limbs of Henle which surround an inner ring of collecting ducts, which in turn surround a central core of thin limbs of Henle. The Savannah Sparrow has a significantly higher volume of medulla compared to the two more mesic species. Within the cortex, the Savannah Sparrow also has a significantly higher volume of proximal tubules but a significantly lower volume of distal tubules than the other species. Within the medulla, the Savannah Sparrow has a significantly higher volume and surface area of capillaries, and a significantly higher surface area of thick limbs of Henle and collecting ducts than the mesic species. These data suggest that the salt marsh Savannah Sparrow has the renal morphology necessary to produce a more highly concentrated urine than the mesic zone species.     

The Pinus halepensis Mill. forests in the central-eastern European Mediterranean basin

The Pinus halepensis Mill. forest vegetation of the central-eastern European Mediterranean basin (France, Italy, Croatia, Albania, Greece) is described here. This study was carried out considering published and unpublished phytosociological data, with the organisation of a systematic classification (syntaxonomic scheme) of the order Pinetalia halepensis according to the principal floristic variations, at multiple spatial scales and according to the physiognomic–structural, floristic, ecological and biogeographical characteristics. Five alliances and 25 associations are recognised.     

Effects of a nutrient pulse supply on nutrient status of the Mediterranean trees Quercus ilex subsp. ballota and Pinus halepensis on different soils and under different competitive pressure

Nutrient availability is a key factor in Mediterranean ecosystems that affects the primary productivity and the community structure. The great variability of its natural availability is now increasing due to frequent fires, pollution events and changes in rainfall regime associated to climate change. Quercus ilex ssp. ballota and Pinus halepensis are the most abundant tree species in the NW Mediterranean basin. They frequently compete in the early and middle successional stages. We investigated the effects of N and P pulse supplies on nutrient uptake capacity in these two species in an after-fire field area and in nursery conditions on different soil types and competing conditions. In the field, N fertilisation had weak effects on nutrient concentration and mineralomass likely as a consequence of this nutrient not being limiting in this field site whereas P fertilisation increased the P mineralomass and the Mg, S, Fe, K and Ca concentrations and mineralomass in the different biomass fractions of both species 1 and 3 years after fertilisation application. In the nursery experiments, P fertilisation increased the mineralomass and concentrations of P, Mg, S, Fe, K and Ca in all biomass fractions including the roots in both species and in different soils and competition conditions. The increment of nutrient mineralomass was due to both the increase of growth and of nutrient concentrations. Both species were able to absorb significant amounts of the P applied by fertilisation (between 5 and 20%) in short time (18 months). Competing vegetation decreased the positive effects of P fertilisation, and in many cases the negative effect of competing vegetation on nutrient mineralomass was stronger when P availability was increased by fertilisation. Q. ilex subsp ballota showed a greater competitive ability for P than the more pioneer species Pinus halepensis in the field but not in the nursery conditions. Pinus halepensis had greater nutrient mineralomass in calcareous than in siliceous soils. Q. ilex subsp. ballota had a higher root biomass allocation and root nutrient allocation than P. halepensis, but both species showed a high capacity to increase their nutrient uptake when its availability increased by fertilisation, thus assuring a great nutrient reserve for future growth periods and contributing to retain nutrients in the ecosystem.     

Plastic bimodal growth in a Mediterranean mixed-forest of Quercus ilex and Pinus halepensis

Mediterranean tree species have evolved to face seasonal water shortages, but may fail to cope with future increases in drought frequency and intensity. We investigated stem radial increment dynamics in two typical Mediterranean tree species, Aleppo pine (Pinus halepensis), a drought-avoiding species, and holm oak (Quercus ilex), a drought-tolerant species, in a mixed forest and on contrasting slope aspects (south- and north-facing). Intra- and inter-annual growth patterns were modelled using the VS-Lite2 model for each tree species and slope-aspect. Both species showed a bimodal growth pattern, with peaks coinciding with favourable conditions in spring and autumn. A bimodal growth pattern is always observed in P. halepensis, while in Q. ilex is facultative, which suggests different strategies adopted by these species to cope with summer drought. More specifically, trees on south-facing slope showed a more evident bimodal pattern and more intra-annual density fluctuations. In recent decades, the intensity of both growth peaks has diminished and drifted away due to the increased summer drought. The VS-Lite2 model reveals a niche partitioning between both species. Differences in growing season’s length and timings of growth peaks in both species are relevant for their coexistence and should be considered for estimating mixed-forest responses under climate change scenarios.     

Somatic embryogenesis in Pinus halepensis Mill.: an important ecological species from the Mediterranean forest

Pinus halepensis Mill. is a common forest species in the Mediterranean area and it is important for environmental conservation. This study established a method of regenerating Pinus halepensis Mill. through somatic embryogenesis. The effect of culture medium (mineral salts, nitrogen source and plant growth regulators), collection date and seed family on embryogenic tissue initiation and proliferation in Pinus halepensis was analysed during the first steps of embryogenesis process. This study showed a marked effect of the culture medium tested as well as some significant differences among collection dates. Furthermore, the embryogenic tissue initiation was affected by the amino acid mixture in the culture medium and the proliferation stage was significantly affected by the combination of plant growth regulators. At the end of the maturation phase the presence of activated charcoal was also evaluated. Finally, maturation of embryogenic tissue was affected by the nitrogen source in the culture medium and these results were different for high and low mature embryo producing cell lines. To the best of our knowledge, this is the first report on Aleppo pine somatic embryogenesis describing a simple and efficient procedure for large-scale somatic embryo production.     

Hydraulic properties of Pinus halepensis,Pinus pinea and Tetraclinis articulata in a dune ecosystem of Eastern Spain

The hydraulic properties of Pinus pinea, Pinus halepensis and Tetraclinis articulata were studied in a coastal dune area from Eastern Spain. The measured variables include vulnerability to xylem embolism (vulnerability curves), hydraulic conductivity and carbon isotopic discrimination in leaves. Leaf water potentials were also monitored in the three studied populations during an extremely dry period. Our results showed that roots had always wider vessels and higher hydraulic conductivity than branches. Roots were also more vulnerable to xylem embolism and operated closer to their hydraulic limit (i.e., with narrower safety margins). Although it was not quantified, extensive root mortality was observed in the two pines during the study period, in agreement with the high values of xylem embolism (> 75%) predicted from vulnerability curves and the water potentials measured in the field. T. articulata was much more resistant to embolism than P. pinea and P. halepensis. Since T. articulata experienced also lower water potentials, safety margins from hydraulic failure were only slightly wider in this species than in the pines. Combining species and tissues, high resistance to xylem embolism was associated with low hydraulic conductivity and with high wood density. Both relationships imply a cost of having a resistant xylem. The study outlined very different water-use strategies for T. articulata and the pines. Whereas T. articulata had a conservative strategy that relied on the low vulnerability of its conducting system to drought-induced xylem embolism, the two pines showed regulatory mechanisms at different levels (i.e., embolism, root demography) that constrained the absorption of water when it became scarce. This revised version was published online in August 2006 with corrections to the Cover Date.     

Ecophysiological response to severe drought in Pinus halepensis Mill. trees of two provenances

As a result of predicted regional climatic changes the need to select for the more drought-tolerant genotypes (ecotypes) among Mediterranean conifers has become clear. Aleppo pine (Pinus halepensisMill.) seems to be one of the most drought-tolerant pine species. Nevertheless, the existence of geographical trends in their genetic differentiation indicates potentially large differences in drought-tolerance among provenances. This assumption was verified by the finding of large variation among provenances in their internal water relations. Hence, the aim of this study was to compare the ecophysiological behaviour of several Aleppo pine provenances under contrasting climatic conditions. Growth parameters (height and diameter) and survival rate were measured in two provenance trials, one planted under sub-desertic conditions at the northern edge of the Negev desert, and the second under thermo-Mediterranean climatic conditions in the central coastal plains, Israel. Ecophysiological parameters such as: predawn needle water potential, sap flow in the xylem (i.e. transpiration), photosynthesis and water-use efficiency were measured in trees of selected provenances. The results suggest that it is not possible to predict provenance performance under harsh conditions from their performance under more favourable ones. Therefore, selection must be carried out under the exact conditions in which the trees from the resultant selection will be planted. The present study clearly emphasises the need for broad selection programs of P. halepensis.     

Genetic evidence for a Janzen-Connell recruitment pattern in reproductive offspring of Pinus halepensis trees

Effective seed dispersal, combining both dispersal and postdispersal (establishment) processes, determines population dynamics and colonization ability in plants. According to the Janzen-Connell (JC) model, high mortality near the mother plant shifts the offspring establishment distribution farther away from the mother plant relative to the seed dispersal distribution. Yet, extending this prediction to the distribution of mature (reproductive) offspring remains a challenge for long-living plants. To address this challenge, we selected an isolated natural Aleppo pine (Pinus halepensis) population in Mt. Pithulim (Israel), which expanded from five ancestor trees in the beginning of the 20th century into ～2000 trees today. Using nine microsatellite markers, we assigned parents to trees established during the early stages of population expansion. To elucidate the effect of the distance from the mother plant on postdispersal survival, we compared the effective seed dispersal kernel, based on the distribution of mother-offspring distances, with the seed dispersal kernel, based on simulations of a mechanistic wind dispersal model. We found that the mode of the effective dispersal kernel is shifted farther away than the mode of the seed dispersal kernel, reflecting increased survival with increasing distance from the mother plant. The parentage analysis demonstrated a highly skewed reproductive success and a strong directionality in effective dispersal corresponding to the wind regime. We thus provide compelling evidence that JC effects act also on offspring that become reproductive and persist as adults for many decades, a key requirement in assessing the role of postdispersal processes in shaping population and community dynamics.     

Climate impact on growth dynamic and intra-annual density fluctuations in Aleppo pine (Pinus halepensis) trees of different crown classes

Mediterranean environments are of special interest for the study of the relationships between climate, growth and anatomic features. Dendrochronological techniques were applied at eight sampling sites that were selected throughout the natural distribution area of Pinus halepensis in the Iberian Peninsula. The objectives of this paper were: (i) to identify relationships between radial growth and climate for different crown classes of Aleppo pine (P. halepensis Mill.); (ii) to quantify the presence of intra-annual density fluctuations (IADFs) according to crown class and cambial age; (iii) to establish the relationships between IADFs and climate. In the more mesic sites, dominant trees showed higher climatic sensitivity than suppressed trees, while in the more xeric sites suppressed trees showed higher sensitivity than dominant trees. Tree-ring growth of both crown classes correlated positively with precipitation during and prior to the growing season. IADFs were more frequent in young than in old stands without differences between crown classes. Precipitation in April and December was positively correlated to the occurrence of IADFs, while precipitation in July correlated negatively. A higher frequency in IADFs occurred in the last 50 years, which coincides with the increase in drought events in the Iberian Peninsula.     

Invasion of Pinus halepensis from plantations into adjacent natural habitats

Questions: Do invasions of P. halepensis (Aleppo pine) from plantations into adjacent natural communities occur in the Mediterranean region, where the species is native? What are the spatio‐temporal processes involved in pine invasions in contrasting Mediterranean and semi‐arid climatic regions? Location: Mediterranean and semi‐arid regions of Israel. Methods: The density of invading Pinus was measured in relation to the distance from the plantation edge. Plants were categorized by age, height, basal stem girth and developmental stage, their spatial distribution was also recorded. Results: Analysis of plant age distribution indicates that the invasion process started when the plantations were 20–25 years old. Most invading plants were found within 20 m from the plantation edge, but a few individuals reached distances up to 100 m and became new invasion foci. Plant density declined sharply with distance from adult trees, data showing a better fit to a power model than to a negative exponential model. Invading Pinus began to produce cones earlier in the semi‐arid than in the Mediterranean region (9 vs 12 years to 50% reproductive plants). In both regions, higher densities of invading plants were found on the west side of the plantation, the opposite direction to the hot winds that prevail during seed release. Conclusion: The frontal advance of P. halepensis from plantations is relatively slow, but the populations also expand by a saltation process, creating spreading ‘islands’ of pine trees in the natural vegetation. Spatial pattern of recruits with distance from the source population was remarkably similar to the pattern of seed dispersal in the same region (Nathan et al. 1999). This implies that the probability of a dispersed seed developing into a plant is independent of the distance from the forest edge.     

Stand structure modulates the long-term vulnerability of Pinus halepensis to climatic drought in a semiarid Mediterranean ecosystem

We investigated whether stand structure modulates the long-term physiological performance and growth of Pinus halepensis Mill. in a semiarid Mediterranean ecosystem. Tree radial growth and carbon and oxygen stable isotope composition of latewood (δ(13)C(LW) and δ(18)O(LW), respectively) from 1967 to 2007 were measured in P. halepensis trees from two sharply contrasting stand types: open woodlands with widely scattered trees versus dense afforested stands. In both stand types, tree radial growth, δ(13)C(LW) and δ(18)O(LW) were strongly correlated with annual rainfall, thus indicating that tree performance in this semiarid environment is largely determined by inter-annual changes in water availability. However, trees in dense afforested stands showed consistently higher δ(18)O(LW) and similar δ(13)C(LW) values compared with those in neighbouring open woodlands, indicating lower stomatal conductance and photosynthesis rates in the former, but little difference in water use efficiency between stand types. Trees in dense afforested stands were more water stressed and showed lower radial growth, overall suggesting greater vulnerability to drought and climate aridification compared with trees in open woodlands. In this semiarid ecosystem, the negative impacts of intense inter-tree competition for water on P. halepensis performance clearly outweigh potential benefits derived from enhanced infiltration and reduced run-off losses in dense afforested stands.     

Intracellular Transport and Elimination of Resin from Epithelial Duct-cells of Pinus halepensis

The ultrastructure of the resin secreting cells of root ductsof young Pinus halepensis seedlings was studied. It is suggestedthat the endoplasmic reticulum (ER) in addition to taking partin resin synthesis also plays a role in transporting the resinfrom the plastids, mitochondria and nuclear envelope to theplasmalemma. By fusing with the plasmalemma the ER releasesthe resin to the outside of the protoplast. The resin producedin the ground cytoplasm and by the Golgi apparatus seems tobe eliminated by plasmalemma invaginations. Pinus halepensis, resin secretion, root ducts, endoplasmic reticulum     

Twig morphology and anatomy of Mediterranean trees and shrubs related to drought tolerance

De Micco V. and Aronne G. 2008. Twig morphology and anatomy of Mediterranean trees and shrubs related to drought tolerance. Bot. Helv. 118: 139 – 148. Woody species populating Mediterranean ecosystems have evolved phenological, physiological and morphological adaptations to summer drought, but their degree of drought tolerance varies, so that different species dominate along gradients from xeric to more mesic environments. In this study, we analysed morpho-anatomical properties of leaves and twigs of eight Mediterranean woody species. To test whether there is a consistent pattern of adaptation to drought, we hypothesised that the rank order of species along the xeric-mesic gradient would be reflected by increasing or decreasing trends in twig properties or in relationships between leaf area and branch size. One-year-old twigs were sampled from plants growing in a common site on the Tresino promontory in southern Italy. Measurements included leaf number and area, branch length and diameter, as well as the relative amounts of pith, xylem and cortex tissues at the basis of branches. All traits varied significantly among species, but none of them correlated with the rank order of species along the xeric-mesic gradient. In addition, the relationship between leaf area and branch size (length or diameter) was not consistently related to the drought tolerance of the species. Inconsistencies could partly be explained by variations in twig anatomy. These results support the view that not single traits, but combinations of morpho-anatomical features influence the overall capability of species to withstand dry conditions. Submitted 12 May 2008; Accepted 24 September 2008 Subject editor: Sabine Güsewell     

On the response of pigments and antioxidants of Pinus halepensis seedlings to Mediterranean climatic factors and long-term ozone exposure

An experiment was carried out in open-top chambers located in eastern Spain. One-yr-old Pinus halepensis Mill. seedlings were exposed during three consecutive summers to the following ozone (O₃) treatments: charcoal-filtered air (CFA), non-filtered air (NFA) or non-filtered air plus 40 nl l⁻¹ O₃, 9 h d⁻¹, 5 d wk⁻¹ (NFA+40). Seasonal variations in Aleppo pine performance were observed since reductions in chlorophyll and cellular peroxidase levels associated with increases in superoxide dismutase activity, were recorded during the summer. Similarly, a reduction in epoxidation state was found at midday during the summer, derived from an activation of the xanthophyll cycle associated to an increment in radiation and temperature levels.
The first O₃-induced effects were recorded in previous-year needles (1991) during the first summer exposure as an increase in extracellular and total peroxidase activities and in zeaxanthin levels in the NFA+40 treatment along with a trend to a higher SOD activity in this treatment. A carry-over effect was detected since a lower winter recovery of chlorophyll levels was found in the NFA+40 seedlings along with a reduction of xanthophyll levels. A reduction in chlorophyll levels was observed in the previous-year needles (1992) from the NFA+40 treatment at the end of the second fumigation period. Realistic ozone exposures induced alterations in plant antioxidative systems and plant pigments as shown in this paper. These observations together with the reductions in stomatal conductance and net photosynthesis recorded in the same experiment, indicate that Aleppo pine is a species sensitive to ozone.