Linking reliance on deep soil water to resource economy strategies and abundance among coexisting understorey shrub species in subtropical pine plantations.

Strategies for deep soil water acquisition (WAdeep ) are critical to a species' adaptation to drought. However, it is unknown how WAdeep determines the abundance and resource economy strategies of understorey shrub species. With data from 13 understorey shrub species in subtropical coniferous plantations, we investigated associations between the magnitude of WAdeep , the seasonal plasticity of WAdeep , midday leaf water potential (Ψmd ), species abundance and resource economic traits across organs. Higher capacity for WAdeep was associated with higher intrinsic water use efficiency, but was not necessary for maintaining higher Ψmd in the dry season nor was it an ubiquitous trait possessed by the most common shrub species. Species with higher seasonal plasticity of WAdeep had lower wood density, indicating that fast species had higher plasticity in deep soil resource acquisition. However, the magnitude and plasticity of WAdeep were not related to shallow fine root economy traits, suggesting independent dimensions of soil resource acquisition between deep and shallow soil. Our results provide new insights into the mechanisms through which the magnitude and plasticity of WAdeep interact with shallow soil and aboveground resource acquisition traits to integrate the whole-plant economic spectrum and, thus, community assembly processes.

[Effects of nitrogen addition on biomass allocation of Pinus koraiensis and Fraxinus mandshurica seedlings under water stress.] / æ°®æ·»åŠ å’Œæ°´åˆ†èƒè¿«å¯¹çº¢æ¾ã€æ°´æ›²æŸ³å¹¼è‹—ç”Ÿç‰©é‡åˆ†é çš„å½±å“.

Nitrogen deposition and climate warming-drying are the main environmental changes in Northeast China. How they affect forest ecosystems has always been studied in ecological research. In this study, we explored the effects of water stress and nitrogen (N) addition on the short-term (in 55 weeks) growth of seedlings of Pinus koraiensis and Fraxinus mandshurica, two key species in broad-leaved and P. koraiensis mixed forests in temperate zone of Northeast China. Results showed that the responses of seedling growth of P. koraiensis and F. mandshurica to nitrogen addition and water stress were significantly different. P. koraiensis was more sensitive to water stress. In the early stage (in 10 weeks) of water stress, leaf biomass of P. koraiensis significantly decreased while the root biomass increased. Nitrogen addition significantly reduced the root and total biomass of P. koraiensis under water stress. F. mandshurica was more sensitive to N addition. Nitrogen addition rapidly increased the stem, root and total biomass of F. mandshurica. Only sustained water stress could significantly affect the stem, root and total biomass of F. mandshurica. Under continuous water stress and N addition, the biomass contribution of leaves and roots to whole seedling, and the ratio of aboveground to underground biomass of both species tended to be constant, indicating strong self-regulation ability of both species. The results suggested that when drought occurred in the future, P. koraiensis would adopt a "positive" adjustment strategy, while F. mandshurica would adopt a "passive" response. The survival rate and adaptability of P. koraiensis would be higher than F. mandshurica. F. mandshurica would benefit more from N deposition than P. koraiensis. Our results are helpful for predicting the changes of community structure in temperate forest in Northeast China.

Maintenance costs of serotiny in a variably serotinous pine: The role of water supply.

Serotiny is an important adaptation for plants in fire-prone environments. However, different mechanisms also induce the opening of serotinous cones in the absence of fire in variably serotinous species. Xeriscence -cone opening driven by dry and hot conditions- is considered to be mediated only by the external environment, but endogenous factors could also play a significant role. Using the variably serotinous Pinus halepensis as our model species, we determined the effects of cone age and scales density in cone opening, and using in-situ and ex-situ manipulative experiments we investigated the role of water availability in the opening of serotinous cones. We hypothesized that loss of connection between the cones and the branch through the peduncles or the absence of water supply could induce a faster cone opening. Results showed that older cones lost more water and opened at lower temperatures, with no influence of scales density. Both field and chamber manipulative experiments (using paired cones of the same whorl) confirmed that water intake through the peduncles affected significantly the pace of cone opening, such that lack of water supply speeded up cone dehiscence. However, this was true for weakly serotinous provenances-more common in this species-, while highly serotinous provenances were indifferent to this effect in the field test. All our results support that cone serotiny in P. halepensis involves the allocation of water to the cones, which is highly consistent with the previously observed environmental effects. Importantly, the existence of maintenance costs of serotinous cones has strong implications on the effects of climate change in the resilience of natural populations, via modifications of the canopy seed banks and recruitment after stand-replacing fires. Moreover, evolutionary models for serotiny in P. halepensis must take into account the significant contribution of maintenance costs to the complex interaction between genotype and the environment.

Differentiation of persistent anatomical defensive structures is costly and determined by nutrient availability and genetic growth-defence constraints.

Conifers exhibit a number of chemical and anatomical mechanisms to defend against pests and pathogens. Theory predicts an increased investment in plant defences under limited nutrient availability, but while this has been demonstrated for chemical defences, it has rarely been shown for anatomical defensive structures. In a long-lived woody plant, we tested the hypothesis that limited nutrient availability may promote an improved differentiation of persistent anatomical defences. We also hypothesized that the costs of differentiation of those long-term anatomical structures may be determined by genetic constraints on early growth potential. Using Pinus pinaster Ait. juveniles, we performed a greenhouse study with 15 half-sib families subjected to experimental manipulation of phosphorus (P) availability and herbivory-related induced responses. When plants were ∼30 cm high, half of the plant material was treated with methyl jasmonate to induce defences, and 2 weeks later plants were harvested and the abundance of resin canals in the cortex and xylem was assessed. Density of constitutive resin canals in the cortex and the total canal system was ∼1.5-fold higher in plants under limited P availability than in fully fertilized plants. Availability of P did not significantly influence the inducibility of resin canal traits. We found negative genetic correlations between plant growth and the density of constitutive canals in the xylem and total canal system, but only under conditions of limited nutrition. These results demonstrate for the first time that differentiation of constitutive anatomical-based defences is affected by P limitation. Moreover, results also evidence the existence of genetic constraints between plant growth and constitutive defensive investment, where lineages with the highest growth potential showed the lowest investment in constitutive resin canals.

Health and vitality assessment of two common pine species in the context of climate change in southern Europe.

The Mediterranean Basin is expected to be more strongly affected by ongoing climate change than most other regions of the earth. The South-eastern France can be considered as case study for assessing global change impacts on forests. Based on non-parametric statistical tests, the climatic parameters (temperature, relative humidity, rainfall, global radiation) and forest-response indicators (crown defoliation, discoloration and visible foliar ozone injury) of two pine species (Pinus halepensis and Pinus cembra) were analyzed. In the last 20 years, the trend analyses reveal a clear hotter and drier climate along the coastline and slightly rainier inland. In the current climate change context, a reduction in ground-level ozone (O3) was found at remote sites and the visible foliar O3 injury decreased while deterioration of the crown conditions was observed likely due to a drier and warmer climate. Clearly, if such climatic and ecological changes are now being detected when the climate, in South-eastern France, has warmed in the last 20 years (+0.46-1.08°C), it can be expected that many more impacts on tree species will occur in response to predicted temperature changes by 2100 (+1.95-4.59°C). Climate change is projected to reduce the benefits of O3 precursor emissions controls leading to a higher O3 uptake. However, the drier and warmer climate should induce a soil drought leading to a lower O3 uptake. These two effects, acting together in an opposite way, could mitigate the harmful impacts of O3 on forests. The development of coordinated emission abatement strategies is useful to reduce both climate change and O3 pollution. Climate change will create additional challenges for forest management with substantial socio-economic and biological diversity impacts. However, the development of future sustainable and adaptive forest management strategies has the potential to reduce the vulnerability of forest species to climate change.

Tradeoffs between soil, water, and carbon -- a national scale analysis from New Zealand.

The tradeoffs between the regulation of soil erosion, provision of fresh water, and climate regulation associated with new Pinus radiata forests in New Zealand are explored using national models. These three ecosystem services for which there is strong demand are monetised as commodities (avoided soil erosion is NZ $1 per tonne; water is NZ $1 per cubic metre; and sequestered carbon is assumed to be NZ $73 per tonne). This permits their summation on a spatial basis to produce a national map of the net benefit of these ecosystem services. Net benefit is spatially variable depending primarily on the relative mix of forest growth rates and demand for irrigation water. New P. radiata forests (once mature) generally reduce mass-movement erosion by an order of magnitude. This provides significant benefits for erosion control where there are high natural rates of erosion. Benefits are especially large in catchments where high sedimentation is increasing flood risk and degrading aquatic ecosystems. The generally high growth rates of P. radiata in New Zealand (8.5 tonnesCha(-1)yr(-1) on average for existing forest) add significant environmental benefits of carbon sinks to climate regulation. However, the reduction of water yield associated with new forests (between 30% and 50%) can neutralise these benefits in catchments where there is demand for irrigation water, such as the eastern foothills of the Southern Alps and the tussock grasslands in the South Island.

Are pine plantations valid tools for restoring Mediterranean forests? An assessment along abiotic and biotic gradients.

The ecological impacts of forest plantations are a focus of intense debate, from studies that consider plantations as "biological deserts" to studies showing positive effects on plant diversity and dynamics. This lack of consensus might be influenced by the scarcity of studies that examine how the ecological characteristics of plantations vary along abiotic and biotic gradients. Here we conducted a large-scale assessment of plant regeneration and diversity in plantations of southern Spain. Tree seedling and sapling density, plant species richness, and Shannon's (H') diversity index were analyzed in 442 pine plantation plots covering a wide gradient of climatic conditions, stand density, and distance to natural forests that act as seed sources. Pronounced variation in regeneration and diversity was found in plantation understories along the gradients explored. Low- to mid-altitude plantations showed a diverse and abundant seedling bank dominated by Quercus ilex, whereas high-altitude plantations showed a virtually monospecific seeding bank of Pinus sylvestris. Regeneration was null in plantations with stand densities exceeding 1500 pines/ha. Moderate plantation densities (500-1000 pines/ha) promoted recruitment in comparison to low or null canopy cover, suggesting the existence of facilitative interactions. Quercus ilex recruitment diminished exponentially with distance to the nearest Q. ilex forest. Richness and H' index values showed a hump-shaped distribution along the altitudinal and radiation gradients and decreased monotonically along the stand density gradient. From a management perspective, different strategies will be necessary depending on where a plantation lies along the gradients explored. Active management will be required in high-density plantations with arrested succession and low diversity. Thinning could redirect plantations toward more natural densities where facilitation predominates. Passive management might be recommended for low- to moderate-density plantations with active successional dynamics (e.g., toward oak or pine-oak forests at low to mid altitudes). Enrichment planting will be required to overcome seed limitation, especially in plantations far from natural forests. We conclude that plantations should be perceived as dynamic systems where successional trajectories and diversity levels are determined by abiotic constraints, complex balances of competitive and facilitative interactions, the spatial configuration of native seed sources, and species life-history traits.

Ecological costs on local adaptation of an insect herbivore imposed by host plants and enemies.

Herbivore populations may become adapted to the defenses of their local hosts, but the traits that maximize host exploitation may also carry ecological costs. We investigated the patterns and costs of local adaptation in the pine processionary moth, Thaumetopoea pityocampa, to its host plants, Pinus nigra and P. sylvestris. The two hosts differ in needle toughness, a major feeding impediment for leaf-eating insects. We observed a west-to-east gradient of increasing progeny size in the Italian Alps, matching the pattern in toughness of their respective local host plant. Eastern populations that feed on the native P. nigra with tough needles had larger eggs, and neonate larvae with larger head capsules, than western populations that feed on the native P. sylvestris and the introduced P. nigra with softer foliage. In a reciprocal transfer experiment that involved the eastern-most and the western-most populations of T. pityocampa from this region, and excluded natural enemies, we found evidence for local adaptation to the host plant. Specifically, larvae from the western population only performed well when raised on their local hosts with soft needles, and they suffered near-complete mortality on the tough foliage at the eastern site. In contrast, larvae from the eastern population survived equally well at both sites. Local adaptation involved a trade-off between progeny size and the number of offspring. We hypothesized that an additional cost, imposed by natural enemies, may be associated with increased egg size: we also observed a west-to-east gradient of increased egg parasitism. We tested this hypothesis in a common garden by exposing eggs of both populations to parasitism by two native egg parasitoids, Ooencyrtus pityocampae and Baryscapus servadeii. The eastern population suffered a higher level of parasitoid attack by O. pityocampae than the western population, and performance of hatched adults of both parasitoids was enhanced in large eggs. Thus, increased neonate quality (larger eggs yielding larger larvae) confers an advantage on tough foliage but incurs the ecological cost of increased parasitism, which may constrain further adaptation by this herbivore.

Component carbon fluxes and their contribution to ecosystem carbon exchange in a pine forest: an assessment based on eddy covariance measurements and an integrated model.

We used a combination of eddy flux, canopy, soil and environmental measurements with an integrated biophysical model to analyze the seasonality of component carbon (C) fluxes and their contribution to ecosystem C exchange in a 50-year-old Scots pine forest (Pinus sylvestris L.) in eastern Finland (62 degrees 47' N, 30 degrees 58' E) over three climatically contrasting years (2000-2002). Eddy flux measurements showed that the growing Scots pine forest was a sink for CO2, with annual net C uptakes of 131, 210 and 258 g C m-2> year-1 in 2000, 2001 and 2002, respectively. The integrated process model reproduced the annual course of daily C flux above the forest canopy as measured by the eddy covariance method once the site-specific component parameters were estimated. The model explained 72, 66 and 68% of the variation in daily net C flux in 2000, 2001 and 2002, respectively. Modeled annual C loss by respiration was 565, 629 and 640 g C m-2 year-1, accounting for 77, 77 and 65% of annual gross C uptake, respectively. Carbon fluxes from the forest floor were the dominant contributors to forest ecosystem respiration, with the fractions of annual respiration from the forest floor, foliage and wood being 46-62, 27-44 and 9-10%, respectively. The wide range in daily net C uptake during the growing season was largely attributable to day-to-day fluctuations in incident quantum irradiance. During just a few days in early spring and late autumn, ecosystem net C exchange varied between source and sink as a result of large daily changes in temperature. The forest showed a greater reduction in gross C uptake by photosynthesis than in C loss by respiration during the dry summer of 2000, indicating that interannual variability in ecosystem net C uptake at this site was modified mostly by summer rainfall and vapor pressure deficit.

[Skewness and inequality of height distribution of young Pinus sylvestris var. mongolica stands introduced on sandy soil with lower limited precipitation for tree survival and normal growth].

Based on the investigation of a transect (20 m wide and 80 m long) along a gradient (bottom, middle, and top) of a dune, whose height was about 5-10 m, the population dynamics in young Pinus sylvestis var. mongolica stands was studied. The results showed that both skewness and height inequality declined before the self-thinning process occurred. Age, initial growth status, density of the population, and neighborhood competition were the main factors affected height inequality. The location at sand dune was also an important factor, and its effect was more obvious than that of density.

Agrobacterium-mediated transformation and assessment of factors influencing transgene expression in loblolly pine (Pinus taeda L.).

This investigation reports a protocol for transfer and expression of foreign chimeric genes in loblolly pine (Pinus taeda L.). Transformation was achieved by co-cultivation of mature zygotic embryos with Agrobacterium tumefaciens strain LBA4404 which harbored a binary vector (pBI121) including genes for beta-glucuronidase (GUS) and neomycin phosphotransferase (NPTII). Factors influencing transgene expression including seed sources of loblolly pine, concentration of bacteria, and the wounding procedures of target explants were investigated. The expression of foreign gene was confirmed by the ability of mature zygotic embryos to produce calli in the presence of kanamycin, by histochemical assays of GUS activity, by PCR analysis, and by Southern blot. The successful expression of the GUS gene in different families of loblolly pine suggests that this transformation system is probably useful for the production of the genetically modified conifers.