Water relations of seedlings of three Quercus species: variations across and within species grown in contrasting light and water regimes

We compared seedling water relations of three Mediterranean Quercus species (the evergreen shrub Q. coccifera L., the evergreen tree Q. ilex L. subsp. ballota (Desf.) Samp. and the deciduous or marcescent tree Q. faginea L.). We also explored seedling potential for acclimation to contrasting growing conditions. In March, 1-year-old seedlings of the three species were planted in pots and grown outdoors in a factorial combination of two irrigation regimes (daily (HW) and alternate day watering (LW)) and two irradiances (43 and 100% of full sunlight). At the end of July, predawn and midday water potentials (Psi(pd), Psi(md)) were measured, and pressure-volume (P-V) curves were obtained for mature current-year shoots. Species exhibited similar Psi(pd) and Psi(md) values, but differed in leaf morphology and water relations. The evergreens possessed larger leaf mass per area (LMA) and were able to maintain positive turgor pressure at lower water potentials than the deciduous species because of their lower osmotic potential at full turgor. However, the three species had similar relative water contents at the turgor loss point because Q. faginea compensated for its higher osmotic potential with greater cell wall elasticity. Values of Psi(pd) had a mean of -1.12 MPa in LW and -0.63 MPa in HW, and Psi(md) had a mean of -1.13 MPa in full sunlight and -1.64 MPa in shade, where seedlings exhibited lower LMA. However, the P-V curve traits were unaffected by the treatments. Our results suggest that Q. faginea seedlings combine the water-use characteristics of mesic deciduous oak and the drought-tolerance of xeric evergreen oak. The ability of Q. coccifera to colonize drier sites than Q. ilex was not a result of higher drought tolerance, but rather may be associated with other dehydration postponement mechanisms including drought-induced leaf shedding. The lack of treatment effects may reflect a relatively low contrast between treatment regimes, or a low inherent responsiveness of these traits in the study species, or both.     

Growth and phenology of seedlings of four contrasting slash pine families in ten nitrogen regimes

Seedlings of two fast- and two slow-growing families of slash pine, Pinus elliottii Englm. var. elliottii, were grown in a greenhouse for one growing season in one of 10 nitrogen (N) regimes. Increasing the N concentration in the nutrient solution resulted in both increased growth rates during the exponential growth phase and extended duration of the growing season. The two components of total height, free growth (epicotyl length to the first bud) and summer growth (height growth after the first bud), both increased significantly with increasing N concentrations up to 40-60 mg l(-1) but decreased at N concentrations above 180 mg l(-1). Compared to seedlings grown in the presence of an optimum N concentration, seedlings grown in the presence of trace amounts of N were smaller and had less summer growth as a percentage of total growth, earlier cessation of height growth, fewer flushes, lower shoot/root ratio, higher root fibrosity, and lower N concentrations in all seedling tissues. Compared to slow-growing families, fast-growing families had more summer height growth, more flushes and later growth cessation, higher shoot/root ratios and higher root fibrosity at all N concentrations. In the presence of an optimum or higher concentration of N, the fast-growing families also had higher needle and total N concentrations than the slow-growing families. Strong family by N-treatment interactions occurred for height, phenology and biomass traits because of the extra responsiveness of one family to increasing N concentration. Several seedling traits were identified that appear promising for predicting field performance in slash pine. The results indicated that the nutrient environment greatly influences genetic expression (e.g., family patterns of summer growth were most closely related to field rankings for seedlings in the trace-N treatment).     

Water relations of pine seedlings in contrasting overstory environments

Overstory conditions influence understory microclimate and resource availability, leading to gradients in evaporative demand and moisture availability that influence seedling water relations. Partial canopies may either reduce seedling moisture stress by ameliorating environmental conditions, or increase moisture stress by reducing soil moisture availability. This study used stable isotope ratios of oxygen (δ¹⁸O) and carbon (δ¹³C) and mass-based foliar nitrogen concentrations to investigate changes in transpiration (E), stomatal conductance (g_s) and intrinsic water use efficiency (iWUE) of pine seedlings across an overstory gradient from open canopy gap environments to closed canopy forest. Foliar δ¹⁸O increased sharply from basal areas of 0–10 m² ha⁻¹ in Pinus banksiana, Pinus resinosa, and Pinus strobus seedlings, followed by a more gradual increase with further increases in basal area. Foliar δ¹³C followed a similar, but less pronounced pattern in P. banksiana and P. strobus seedlings, and had no apparent relationship with overstory basal area in P. resinosa seedlings. The slope of the δ¹⁸O:δ¹³C relationship was positive for every species. Foliar nitrogen concentrations were not correlated with overstory basal area. These results suggest seedling E declined as overstory basal area increased due to reductions in g_s, while iWUE increased slightly from open gaps to partial canopy environments. Open gap environments appear to provide sufficient moisture to sustain high leaf-level gas exchange rates in the species we studied, while relatively small increases in overstory basal area apparently promote rapid declines in g_s, leading to greatly reduced seedling water loss and small increases in iWUE.     

Freezing injury in primary and secondary needles of Mediterranean pine species of contrasting ecological niches

? Pine seedlings show a marked ontogenetic difference between primary and secondary needles, the latter prevailing with a different timing among species.

? Using the electrolyte leakage method following an artificial freezing test, we aimed at (1) determining the differences in freezing tolerance between primary and secondary needles in eight pines of contrasting thermal habitats: P. canariensis, P. pinea, P. halepensis, P. brutia, P. pinaster, P. nigra, P. sylvestris and P. radiata, (2) evaluating the relation between freezing tolerance and sclerophylly and (3) relating freezing tolerance with the climate of origin.

? Primary needles were significantly more sensitive to freezing than secondary needles in Pinus halepensis, P. brutia, P. pinaster and P. nigra, whereas no differences were found in P. canariensis, P. pinea and P. radiata. LT50 was uncorrelated with needle sclerophylly but very highly correlated with the mean temperature of the coldest month at the seed source.

? Results support an adaptive role of secondary needles in the mountain Mediterranean pines P. nigra, P. pinaster and P. brutia, while the more complex responses in coastal Mediterranean pines can be interpreted in the light of seedling ontogeny and species’ ecological niches.

     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

In vivo and in situ rhizosphere respiration in Acer saccharum and Betula alleghaniensis seedlings grown in contrasting light regimes

A perfusive method combined with an open-system carbon dioxide measurement system was used to assess rhizosphere respiration of Acer saccharum Marsh. (sugar maple) and Betula alleghaniensis Britton (yellow birch) seedlings grown in 8-l pots filled with coarse sand. We compared in vivo and in situ rhizosphere respiration between species, among light regimes (40, 17 and 6% of full daylight) and at different times during the day. To compute specific rhizosphere respiration, temperature corrections were made with either species-specific coefficients (Q10) based on the observed change in respiration rate between 15 and 21 degrees C or an arbitrarily assigned Q10 of 2. Estimated, species-specific Q10 values were 3.0 and 3.4 for A. saccharum and B. alleghaniensis, respectively, and did not vary with light regime. Using either method of temperature correction, specific rhizosphere respiration did not differ either between A. saccharum and B. alleghaniensis, or among light regimes except in A. saccharum at 6% of full daylight. At this irradiance, seedlings were smaller than in the other light treatments, with a larger fine root fraction of total root dry mass, resulting in higher respiration rates. Specific rhizosphere respiration was significantly higher during the afternoon than at other times of day when temperature-corrected on the basis of an arbitrary Q10 of 2, suggesting the possibility of diurnal variation in a temperature-independent component of rhizosphere respiration.     

Carbon allocation and morphology of cherrybark oak seedlings and sprouts under three light regimes

• ? Continued problems in regenerating oak forests has led to a need for more basic information on oak seedling biology.
• ? In the present study, carbon allocation and morphology were compared between cherrybark oak (Quercus pagoda Raf.) seedlings and sprouts at 1-Lag grown in full, 47%, and 20% sunlight.
• ? Results indicated that cherrybark oak seedling carbon allocation and morphology responded plastically to light availability. In full light, roots were sinks for ¹⁴C, while shoots were sinks for ¹⁴C under reduced light availability. Cherrybark oak sprouts exhibited similar carbon allocation patterns in response to light availability, but displayed stronger shoot sinks than seedlings when grown underreduced light availability. We also showed that young oak sprout roots are a sink for ¹⁴C-photosynthates.
• ? Results from this study point to the need for a morphological index for oak sprout development so more precise comparisons in sprout development and physiology can be made with seedlings.

     

Effects of contrasting fertilization and moisture regimes on biomass,nutrients, and water relations of container grown red pine seedlings

Red pine seedlings were grown for 16 weeks under contrasting fertilizat (conventional, exponential) and moisture (wet, moist, dry) regimes to assess preconditioning effects of treatments on biomass production, nutrient uptake and allocation, and water relations. Growth, nutrient status, and water relations were affected more by moisture availability than by fertilization regime. Exponential fertilization under limited irrigation lowered shoot/root mass ratio, increased root nutrient reserves, and enhanced drought avoidance compared to conventional fertilization regimes. Drought treatments decreased nutrient uptake in the shoots of both fertilization regimes by 24%, but increased nutrient accumulation in the roots by 39% in the exponential regime compared to 17% in the conventional. These results may explain improved outplanting performance noted for exponentially fertilized container stock.     

Divergence among species and populations of Mediterranean pines in biomass allocation of seedlings grown under two watering regimes

Seedlings of four populations each of Pinus pinaster, P. halepensis, P. canariensis and P. pinea were grown in controlled conditions to evaluate both inter- and intra-specific differences in response to watering. We submitted half of the plants to a moderate water stress and after 22 weeks, we recorded height, stem diameter and root, stem and leaves dry weight. Patterns and amounts of phenotypic changes, including changes in biomass allocation, were analysed. We found a scant response in P. canariensis, P. pinaster and P. halepensis presented high population divergence for phenotypic changes, and P. pinea showed marked allocational shifts and no population divergence. The phenotypic changes observed within species are interpreted as a plastic response. The variation encountered within P. halepensis and P. pinaster may be indicative of specialisation to either resource-rich or resource-poor habitats, being populations from favourable sites more plastic. P. pinea exhibited a very uniform plastic response, indicating generalist behaviour.     

Performance of ponderosa pine seedlings under four herbicide regimes in northeast Oregon

The herbicide, hexazinone, was applied four ways over ponderosa pine, 2–0 seedlings planted in northeast Oregon. The four treatments were two broadcast applications, a single broadcast application, a large spot application, and a small spot application. Seedling survival and growth were monitored for five growing seasons. Results indicate that survival more than doubled with either large or small spot applications compared to no application, and either one or two broadcast applications can increase survival an additional 30% over spot applications. Differences in stem volume were substantial, with two broadcast applications yielding more than twice the volume of a single broadcast treatment and more than five times the volume of seedlings treated with spot applications. Trees in small spots were still three times bigger than surviving seedlings in the control. These results are consistent with the concept of competition threshold. Management implications were considered in terms of cost of established seedlings. Although broadcast applications cost more per acre than spot applications, gains in seedling survival, growth and quality offset the additional cost and translate into lower established seedling costs. The cost effectiveness of broadcast applications also may be seen in the elimination of replanting or in-planting requirements and increases in long-term growth potential of the established trees.     

Spatio-temporal perspectives of forest fires regimes in a maturing Mediterranean mixed pine landscape

It has been suggested that during the past several decades, the frequency and the intensity of wildfires have markedly increased in the Mediterranean basin. We came to assess this postulation in the forested region of Mount Carmel, Israel. This region is characterized by Quercus spp. and Pistacia spp. maquis and has been intensely afforested with Pinus spp. stands since the 1920s. We compiled a GIS-based database of the fires recorded in the region since 1983, in addition to archiving data beginning from the 1940s. The data were collected from land stewardship agencies’ archives, fire departments and aerial photographs. Prior to the early 1980s, no systematic documentation of the fires was available, rather just sporadic qualitative documentation of the large forest fires that occurred. Between 1944 and 1982, only 6 large fires were documented, while after that 11 large fires occurred. Analysis indicated that the spatial distribution of the fires does not occur at random, and their locations are significantly closer to roadsides compared to an expected random distribution. The annual number of fires and the areas burned during the last two decades were not correlated with annual precipitation in any manner. Accordingly, we suggest that the increased number of large forest fires during the last decades is associated with the maturation and senescence of the planted forest coupled with increased human activities.     

Growth regulation of Scots pine seedlings with different fertilizer compositions and regimes

Scots pine (Pinus sylvestris L.) seedlings were grown in containers filled with peat, using two different fertilizers and three different fertilizer regimes. Seedling shoot and root growth and shoot content of nitrogen, potassium and phosphorus were followed in the nursery and after outplanting in the field. Attempts to regulate growth rate by an exponential nutrient supply were not successful, but the root/shoot ratio was influenced by the fertilization regime. Internal nitrogen concentration was stable only for seedlings with low relative growth rate, while seedlings with high nutrient supply in the nursery showed strong nutrient dilution in the shoot after planting.     

Interactive effects of shade and irrigation on the performance of seedlings of three Mediterranean Quercus species

Shade and irrigation are frequently used to increase the success of Mediterranean Quercus spp. plantations. However, there is controversy about the combined effects of these treatments on plant performance. We assessed the effects of two irradiances (full sunlight and moderate shade) and two summer watering regimes (high (daily) and low (alternate days)) on leaf and whole-plant traits of 1-year-old seedlings of Quercus coccifera, Q. ilex subsp. ballota and Q. faginea grown outdoors for 8.5 months. Leaf traits included measures of morphology, nitrogen concentration, gas exchange and photochemical efficiency, and measures of whole-plant traits included biomass allocation patterns, growth phenology, across-summer leaf area change and relative growth rate (RGR). Moderate shade reduced leaf mass per area, increased photochemical efficiency, maximum carbon assimilation rate (Amax) and allocation to leaves, and prolonged the growing period in one or more of the species. Daily watering in summer increased Amax of Q. ilex and prolonged the growing period of Q. ilex and Q. faginea. Both treatments tended to increase RGR. The effect of shade was greater in the low-watering regime than in the high-watering regime for two of the 15 studied traits, with treatment effects being independent for the remaining 13 traits. Leaf nitrogen and the ability to maintain leaf area after the arid period, rather than biomass allocation traits, explained the variation in seedling RGR. Trait responsiveness to the treatments was low and similar among species and between study scales, being unexpectedly low in Q. faginea leaves. This may be because selective pressures on leaf plasticity act differently in deciduous and evergreen species. We conclude that moderate shade and daily summer watering enhance the performance of Mediterranean Quercus seedlings through species-specific mechanisms.     

The role of cold storage and seed source in the germination of three Mediterranean shrub species with contrasting dormancy types

Context

The use of native species in ecological restoration is highly recommended but, in practice, it is often impaired by knowledge gaps in the germination ecology of suitable species.

Aims

This study aimed to assess the role of storage conditions and seed source on the germination of three Mediterranean shrub species with contrasting types of dormancy.

Methods

Ripe fruits were harvested at two or three distant locations in mainland Portugal. Seeds were subjected to three treatments consisting in different storage conditions: cold storage at low and high moisture conditions, plus a control. Five replicates of up to 30 seeds were placed under constant temperature conditions and germination was monitored weekly during 14 weeks.

Results

The effect of cold storage at high moisture on germination differed between the three species and seed source played a significant role in the germination of all three species. In the case of the species with dormancy, the observed differences in germination could reflect changes in the species’ dormancy degree or sensitivity to dormancy breaking factors across their geographical range. In the case of Pistacia lentiscus (no dormancy), the results suggested a possible adaptation of the northern seed source to high moisture conditions.

Conclusions

The observed differences between species agreed well with their dormancy types, and the seed source-related differences could be adaptive features, as they seemed related with local climate conditions.     

Early growth and photosynthetic responses to light in seedlings of three tropical species differing in successional strategies

Containerized seedlings of three commercially important tropical species were grown under four different light treatments [i.e., 100 (open site), 45, 22 and 10% sunlight] for 130 days. Light-saturated photosynthesis (A _max) and light saturation estimates (LSE) reflected the species successional status with Terminalia superba Engl. and Diels, the pioneer species showing largest mean A _max and LSE at 100% sunlight, whereas at 10% sunlight, it showed the lowest A _max and LSE. At 22% sunlight, Cedrela odorata L., an intermediate successional species had greater A _max and LSE than Mansonia altissima A. Chev., a non-pioneer light demander and T. superba. T. superba had the lowest relative growth rate (RGR) at 10% sunlight and greatest net assimilation rate (NAR) at 100% sunlight; although a higher RGR at this light level was not seen for this species. Strong and positive linear mean A _max–mean NAR relationship of C. odorata and T. superba indicated that differences in leaf photosynthetic rates of the two species were reflected in their NAR, which increased with increasing light. At final harvest, superior biomass production was found at 45% sunlight for all the species. Seedling responses in specific leaf area, leaf area ratio, leaf mass ratio and root mass ratio were typically those found along a light gradient. At the 100% sunlight, intrinsic water-use efficiency (WUE), F _v/F _m and final root system of the plants was generally superior in T. superba but at 10% sunlight, WUE was inferior in T. superba when compared to C. odorata and M. altissima, reflecting the respective species’ short-term acclimation to high or low light. Results of this study may have practical use in screening tropical tree species for use in plantation forestry.     

Water stress responses of seedlings of four Mediterranean oak species

Effects of water stress on phenology, growth, stomatal activity and water status were assessed from April to November 1996 in 2-year-old seedlings of Quercus frainetto Ten. (Quercus conferta Kit.), Quercus pubescens Willd., Quercus macrolepis Kotschy (Quercus aegilops auct.) and Quercus ilex L. growing in containers in northern Greece. All four species developed more than 50% of their total leaf area before the beginning of June--an adaptation to arid climates. Well-irrigated plants tended to develop greater individual leaf area, number of leaves per plant, total plant leaf area, height and root:shoot ratios than water-stressed plants, but the difference between treatments was not significant for any parameter in any species. Quercus macrolepis appeared to be the most drought-tolerant of the four species. It maintained the highest number of leaves of the smallest size and increased the proportion of fine roots during drought. In all species, drought caused significant decreases in stomatal conductance and predawn and midday water potentials from mid-July until the end of August, when the lowest soil water content and highest mean daily air temperatures and midday leaf temperatures occurred; however, the responses were species-specific. Among the four species, Quercus macrolepis sustained the highest stomatal conductance despite very low water potentials, thus overcoming drought by means of desiccation tolerance. Quercus ilex decreased stomatal conductance even before severe water stress occurred, thereby avoiding desication during drought. Quercus pubescens had the highest water potential despite a high stomatal conductance, indicating that its leaf water status was independent of stomatal activity. Quercus frainetto was the least drought-resistant of the four species. During drought it developed very low water potentials despite markedly reduced stomatal aperture.     

Photosynthetic light response of flooded cherrybark oak (Quercus pagoda) seedlings grown in two light regimes

Two-year-old cherrybark oak (Quercus pagoda Raf.) seedlings raised in full or partial (27%) sunlight were flooded for 30 days to study the effects of light availability and root inundation on photosynthetic light response. Compared with seedlings receiving full sunlight, seedlings receiving partial sunlight developed leaves with 90% greater blade area, 26% less mass per unit volume, and 35% lower nitrogen (N) concentration per unit area, leading to a 15% reduction in leaf photosynthetic capacity when carbon exchange rates were based on blade area. However, when carbon exchange rates were based on leaf mass, leaves acclimated to partial sunlight exhibited a 15% greater photosynthetic capacity realized primarily through an increased initial slope of the photosynthetic light response (A/PPFD) curve and increased net photosynthesis at leaf saturation (Amax). Short-term flooding increased leaf mass per unit area more than 19%, reduced foliar N concentrations per unit dry mass by 19%, and initiated reductions in Amax and apparent quantum yield (phi) of seedlings in both light regimes. Greatest impairment of Amax (56% area basis, 65% mass basis) and phi (40%) were observed in leaves receiving full sunlight, and the declines were concomitant with a 35% decrease in chlorophyll concentration. Flooding also depressed instantaneous photosynthetic N-use efficiency (PPNUE) such that Amax decreased 54%, and the initial slope of PPNUE/PPFD curves decreased 33 and 50% for leaves acclimated to partial and full sunlight, respectively. The A/PPFD patterns indicated that the magnitude of flood-induced inhibition of the photosynthetic mechanism of cherrybark oak seedlings is determined partly by the light environment.     

Feedbacks between fuel reduction and landscape homogenisation determine fire regimes in three Mediterranean areas

In densely populated areas like the Mediterranean, wildfire extent is mostly limited by fire suppression and fuel fragmentation. Fire is known to spread more easily through high fuel loads and homogenous terrain and it is supposed to reduce fuel amount and continuity, creating a negative feedback. Here we combine information from administration fire records, satellite imagery fire scars and land use/cover maps to asses the effects of fire on landscape structure and vice versa for three areas in Catalonia (NE Spain). We worked with three spatial focuses: the actual fire scar, 1 km² squares and 10 km² squares. In these regions agriculture land abandonment has lead to increased fuel continuity, paralleled by an increment of fire size. We confirm that fire spread is facilitated by land use/cover types with high fuel load and by homogeneous terrain and that fire reduces fuel load by transforming forests into shrublands. But we also found that fire increased landscape homogeneity, creating a positive feedback on fire propagation. We argue that this is possible in landscapes with finer grain than fire alone would create. The lack of discontinuities in the fuel bed diminishes the extinction capacity of fire brigades and increases the risk of large fires. We recommend that fire management should focus more on conservation of the traditional rural mosaic in order to prevent further increases in fuel continuity and fire risk.     

Dependence of needle architecture and chemical composition on canopy light availability in three North American Pinus species with contrasting needle length

Morphology and chemical composition of needles of shade-intolerant southern conifers (Pinus palustris Mill. (mean needle length +/- SD = 29.1 +/- 4.1 cm), P. taeda L. (12.3 +/- 2.9 cm) and P. virginiana Mill. (5.1 +/- 0.8 cm)) were studied to test the hypothesis that foliage acclimation potential to canopy light gradients is generally low for shade-intolerant species, and in particular, because of mechanical limitations, in species with longer needles. Plasticity for each needle variable was defined as the slope of the foliar characteristic versus irradiance relationship. A novel geometrical model for needle area and volume calculation was employed for the three-needled species P. palustris and P. taeda. Needle thickness (T) strongly increased, but width (W) was less variable with increasing daily integrated quantum flux density averaged over the season (Q(int)), resulting in changes in cross-sectional needle shape that were manifested in a positive relationship between the total to projected needle area ratio (A(T)/A(P)) and Q(int) in the three-needled species. In contrast, cross-sectional needle geometry was only slightly modified by irradiance in the two-needled conifer P. virginiana. Needle dry mass per unit total needle area (M(T)) was positively related to Q(int) in all species, leading to greater foliar nitrogen contents per unit area at higher irradiances. Separate examination of the components of M(T) (density (D) and the volume (V) to A(T) ratio; M(T) = DV/A(T)) indicated that the positive effect of light on M(T) resulted solely from increases in V/A(T), i.e., from increases in the thickness of foliage elements. Foliar chlorophyll content per unit mass increased with increasing Q(int), allowing an improvement in light-harvesting efficiency in low light. The variables characterizing needle material properties (D, the dry to fresh mass ratio, and needle carbon content per unit mass) were generally independent of Q(int), suggesting that needles were less stiff and had greater tip deflections under their own weight at lower irradiances because of smaller W and T. Comparisons with the literature revealed that plasticity in foliar characteristics tended to be lower in the studied shade- intolerant species than in shade-tolerant conifers, but plasticity among the investigated species was unaffected by needle length. However, we argue that, because of mechanical limitations, plastic changes in needle cross section in response to low irradiance may decrease rather than increase light-interception efficiency in long-needled species.     

Response to the interaction of thinning and pruning of pine species in Mediterranean mountains

Pruning allows knot-free timber to be obtained, thereby increasing the value of the highest-value wood products. However, the effect of pruning on growth is under discussion, and knowledge about the tree response to the simultaneous development of thinning and pruning is scarce. The objective of this study was to analyze the effect of the interaction of thinning and pruning on tree and stand level and the annual radial growth of two pine species native to Mediterranean mountains. We used long-term data of three trials installed in pine stands where several combinations of pruning and thinning were developed. Five inventories were carried out for each trial, and the mean dasometric features of the different treatments were compared using linear mixed models including a competition index. In addition, we collected cores from ten trees per plot in order to evaluate the annual response of trees to the thinning and pruning. We analyzed the annual radial growth using a semiparametric approach through a smooth penalized spline including rainfall and temperature covariates. Pruning did not show any effect on growth. However, larger diameter and increased annual radial growth were found in thinned plots, both with and without pruning, as compared to unthinned plots. Also, we found significant effects of climate on annual radial growth. We recommend the application of thinning and pruning in stands of Mediterranean mountains in order to get knot-free timber since growth reduction was not found in thinned stands.