Osmotic adjustment in three-year-old seedlings of five provenances of maritime pine (Pinus pinaster) in response to drought

Three-year-old seedlings of five provenances of Pinus pinaster Ait. that differed in climatic conditions at their geographical origin were subjected to decreasing soil water availability. The degree of needle osmotic adjustment (OA) was estimated based on logarithmic plots of needle relative water content (RWC) against needle osmotic potential (Psi(pi)); i.e., lnRWC versus -ln(-Psi(pi)). There were significant differences among provenances in active OA (0.13 to 0.30 MPa for a decrease in RWC to 80%), and a clear negative relationship was found between OA and precipitation (650 to 1280 mm of mean annual rainfall) at the geographical origins of the provenances. A high osmoregulatory capacity contributes to the maintainance of positive turgor at low water potentials. We conclude that OA is one of the mechanisms underlying adaptation to drought in P. pinaster. Solute accumulation was about 2.3 times higher in the provenance from the driest site than in the provenance from the wettest site. The contribution of osmotic adjustment to differences in drought tolerance mechanisms among provenances is discussed.     

Maturation of maritime pine (Pinus pinaster Ait.) seedlings after exposure to a period of continuous light

Nine half-sib families of maritime pine (Pinus pinaster Ait.) of known adult performance were grown in continuous light at either 25 degrees C or 25/20 degrees C for 18 weeks. They were then exposed to a dormancy induction period followed by a dormancy release period and then grown for a further 9 weeks in a 16-h photoperiod at a day/night temperature of 25/20 degrees C. Seedlings exhibited great diversity in morphology at the end of the first growth period. The number of morphogenetic cycles varied between one and three and the form of the apical meristem ranged from a typical rosette to an adult-like bud. The type of seedling obtained at the end of the first growth period strongly influenced later growth, independently of the temperature regime. Maturity was proportional to the number of morphogenetic cycles achieved during the first growth period and was characterized by short growth duration, small primary needles and a high degree of fixed growth. The state of the apical meristem that underwent the dormancy period had less influence on the rate of maturation than the number of morphogenetic cycles. The time course of maturation was endogenously controlled and varied among traits. Conspicuous morphological differences were not associated with changes in the relationship between growth components at the phenotypic level. However, there seemed to be a shift in the genetic correlations between growth components after first budset.     

Growth and water relations of three geographically separate origins of maritime pine (Pinus pinaster) under saline conditions

Water relations and growth of maritime pine (Pinus pinaster Ait.) were investigated in 2-year-old seedlings of French ('Landes'), Iberian ('Iberian') and Moroccan ('Tamjoute') origin raised for 67 days in a flowing solution culture system containing 0, 50, 150 or 250 mM NaCl. Height growth, and stem, needle and root dry matter were reduced by salinity with minor differences among geographic origins. Predawn needle water potential was decreased by salinity and corresponded approximately to the osmotic potential of the nutrient solution. Stomatal conductance was reduced according to the amount of salinity applied. Whole-plant hydraulic conductance was also reduced, even when expressed on a root dry weight basis. The osmotic potential of xylem sap was five- to sixfold lower than that of the nutrient solution. Seedlings of the most southerly origin (Tamjoute) exhibited a greater ability to decrease osmotic potential under saline conditions than seedlings of more northerly origin (Landes and Iberian) as a result of higher mineral cation transport to the shoot.     

Influence of provenance,silvicultural regime and tree shape on the quality of maritime pine (Pinus pinaster) timber

The aim of this study was to establish the sources of variation in maritime pine wood properties related to appearance and strength, because of the importance of this wood in the sawing industry in Galicia (NW Spain). Ten trees from each of ten plots of the species (i.e., 100 trees) growing in Galicia were felled and sawn to produce the structural planks required for studying the presence of external strength-reducing characteristics. The planks were sampled from the basal logs to 16 m height in the stems, yielding cross sections of target dimensions 100 × 40 mm to 200 × 200 mm. We propose a model in which individual tree height and dominant height are used to predict the knottiness of planks from trees in mature stands. The abundance of externally visible knots on faces and edges of sawn planks is therefore more dependent on the stand location than on the prior silvicultural treatment of the stand.     

Variation in growth and osmotic regulation of roots of water-stressed maritime pine (Pinus pinaster Ait.) provenances

Hydroponically cultivated Pinus pinaster Ait. seedlings of a drought-sensitive population from France (Landes) and of a more drought-adapted population from Morocco (Tamjoute) were subjected to a progressive increase in water stress by additions of an osmoticum (polyethylene glycol 600) to the nutrient solution. The final osmotic potential (Psi(ms)) of the nutrient solution was achieved over a period of up to 6 days, and ranged from -0.03 (control, no added osmoticum) to -0.8 MPa. In the 6 days during which water stress was imposed, roots elongated faster in the Moroccan provenance than in the French provenance, but the applied water deficits did not inhibit root elongation in either population. Among treatments, root dry weight per unit root length, total root dry weight and root/shoot dry weight ratio increased with decreasing Psi(ms) in both provenances. Both the water potential (Psi(w)) of the roots (apices) and the water potential difference between the roots and the nutrient solution decreased as Psi(ms) decreased. The reduction in Psi(w) was matched by a decrease of comparable magnitude in cell osmotic potential (Psi(pi)) so that root turgor was unaffected by the Psi(ms). Osmotic adjustment was greater, however, in the Moroccan provenance than in the French provenance. Consequently, under the osmotically imposed water stress, the water potential difference between root and nutrient solution was greater in the Moroccan provenance than in the French provenance. Similar changes in plant water relations were observed when seedlings were grown in drying sand.     

Physical properties and termite durability of maritime pine Pinus pinaster Ait., heat-treated under vacuum pressure

An original heat treatment performed under vacuum pressure was investigated. Maritime pine samples were treated at six different temperatures: 140, 160, 180, 200, 230 and 260°C. The physical and mechanical consequences, i.e. bending strength (MOR), modulus of elasticity (MOE), hygroscopic behaviour, equilibrium moisture contents and anti-swelling efficiency (ASE) were studied. A no-choice feeding test according to the NF EN 117 standard was achieved. Temperatures up to 200°C had no significant effect on wood properties. However, at 230 and 260°C, the decrease in MOR was severe, reaching 42.5 and 62.5%, respectively. Whatever the treatment conditions, wood samples were still highly degraded by termites, revealing no increase in their durability.     

Heat-induced colour changes of pine (Pinus pinaster) and eucalypt (Eucalyptus globulus) wood

Heat treatment of Pinus pinaster and Eucalyptus globulus wood was carried out by hot air in an oven for 2–24 h at 170–200°C and by steam in an autoclave for 2–12 h at 190–210°C. The colour parameters L*, a* and b* were determined by the CIELAB method on radial, tangential and transverse sections of untreated and treated wood, and their variation with regard to the treatment (ΔL*, Δa* and Δb*) were calculated in percent. For untreated eucalypt wood, lightness (L*) varied between 54.1 and 63.8% with a* between 7.4 and 8.5, and b* between 15.7 and 19.9. For untreated pine wood, L* varied between 67.3 and 76.1%, a* between 6.9 and 7.6 and b* between 16.3 and 24.1. Oven heat-treated wood became darker (ΔL* about 50% for 4% mass loss), and this was more for eucalypt wood under the same treatment conditions. In general, the contribution of red (a*) and yellow (b*) colour decreased with heat treatment. The transverse section of the two species darkened less for both the treatments with small differences between radial and tangential sections. Lightness decrease was related to chemical changes; with good correlations with glucose (R ² = 0.96), hemicelluloses (R ² = 0.92) and lignin (R ² = 0.86). As regards colour, the heat treatments showed an interesting potential to improve the wood quality for solid timber products from pine and eucalypt.     

The relationship between seed weight, germination and biochemical reserves of maritime pine (Pinus pinaster Ait.) in Morocco

Seed germination and biochemical reserves of maritime pine (Pinus pinaster Ait.) were studied with the aim of providing germination information for reforestation and conservation programs. Ten natural populations were used to assess variation in seed weight, germination characteristics and biochemical reserves and to examine the relationship between these characteristics. The analysis of variance showed highly significant population effects for seed weight, germination characteristics and protein content in both seeds and the female gametophyte. The mobilization of protein content in female gametophyte during seed germination differed more among populations than sugar content, suggesting that protein content was more sensitive to environment effects than sugar content. A strong positive correlation between germination capacity and the protein content in both seeds and female gametophyte indicated that the best populations in term of germination capacity may also be the richest in protein content. Seeds that were heavier and had a lower speed of protein content mobilization in the female gametophyte appeared to be better adapted to drought conditions. The results also suggested that as much as possible of the potentially valuable genetic variation among populations of this species should be preserved for reforestation and conservation purposes.     

Wood quality assessment of Pinus radiata (radiata pine) saplings by dynamic mechanical analysis

Wood Science and Technology - The use of dynamic mechanical analysis was explored as a possible method of screening for wood quality in breeding programmes. Viscoelastic properties along the grain...     

Coherent response of a group of plantation-grown Scots pine trees to wind loading

Wind-induced motion of 29 neighbouring trees growing in a Scots pine plantation was measured over the period 14 March to 24 March 2008. The bi-orthogonal decomposition (BOD) of the complex tree motion field into a limited number of spatio-temporal modes provided the basis for the analysis of the response behaviour to wind excitation of the group of sample trees. It is shown that the first BOD-mode was the most energetic and differed from all other BOD-modes. The BOD-results as well as the results from wavelet analysis of the temporal eigenvectors of the BOD-modes demonstrate that two concomitant low-frequency components in the streamwise wind velocity component stimulated coherent response of the sample trees at the tree group level. However, in the range of the wind speed measured close to the top of the Scots pine forest canopy (hourly mean wind speed values lower than 6 m s⁻¹), the wind loads associated with these low-frequency airflow structures were too low to harmonise the motion of the sample trees completely. It is shown that instantaneous single tree responses to wind excitation were highly irregular in magnitude and direction. Results from Fourier and wavelet analysis demonstrate that sway in the first mode dominated the wind-induced sway behaviour at the tree level.     

Seasonal and interannual variations in carbon isotope discrimination in a maritime pine (Pinus pinaster) stand assessed from the isotopic composition of cellulose in annual rings

Stable carbon isotope composition (delta; per thousand) was measured on cellulose extracted from maritime pine (Pinus pinaster A?t.) tree rings to investigate inter-tree and interannual variability (7 trees, 20 rings per tree, each ring divided into early and late wood). A model of stand primary production coupled to water balance was used to calculate the stand annual water-use efficiency (WUE). Inter-tree variability in discrimination (maximum 2.88 per thousand in late wood in 1989, 2.69 per thousand in early wood in 1983) was as large as interannual variation (maximum 2.72 per thousand in late wood, 2.05 per thousand in early wood). Tree size did not explain these differences. Relationships were found between annual discrimination and climate variables such as annual rainfall, summer temperature and vapor pressure deficit (VPD). Higher correlations were found with late wood discrimination. Early wood discrimination was shown to be related to previous-year late wood discrimination. Late wood discrimination was also related to soil water availability. Stand annual WUE was only weakly related to tree ring carbon discrimination.     

Crown architecture of understory and open-grown white pine (Pinus strobus L.) saplings

Crown architecture and growth allocation were studied in saplings of eastern white pine (Pinus strobus L.), a species classified as intermediate in shade tolerance. A comparison was made of 15 understory saplings and 15 open-grown saplings that were selected to have comparable heights (mean of 211 cm, range of 180-250 cm). Mean ages of understory and open-grown trees were 25 and 8 years, respectively. Understory trees had a lower degree of apical control, shorter crown length, and more horizontal branch angle, resulting in a broader crown shape than that of open-grown trees. Total leaf area was greater in open-grown saplings than in understory saplings, but the ratio of whole-crown silhouette (projected) leaf area to total leaf area was significantly greater in understory pine (0.154) than in open-grown pine (0.128), indicating that the crown and shoot structure of understory trees exposed a greater percentage of leaf area to direct overhead light. Current-year production of understory white pine was significantly less than that of open-grown white pine, but a higher percentage of current-year production was allocated to foliage in shoots of understory saplings. These modifications in crown structure and allocation between open-grown and understory white pine saplings are similar to those reported for more shade-tolerant fir (Abies) and spruce (Picea) species, but the modifications were generally smaller in white pine. As a result, white pine did not develop the flat-topped "umbrella" crown structure observed in understory fir and spruce, which approaches the idealized monolayer form that maximizes light interception. The overall change to a broader crown shape in understory white pine was qualitatively similar, but much more limited than the changes that occurred in fir and spruce. This may prevent white pine from persisting in understory shade as long as fir and spruce saplings.     

Growth response of Pinus pinaster Ait. to climatic variables in central Spanish forests

? The effect of climatic variables (temperature and precipitation) on radial growth of the Mediterranean Maritime pine (Pinus pinaster Ait.) was studied using dendrochronological techniques in the Iberian Peninsula.

? Ten tree-ring width chronologies, along the central distribution area of the species, were built. Chronology variability was analysed using Principal Component Analysis (PCA) for the period 1952–2005.

? The first principal component (PC1) explained 56% of tree-growth variability. Tree-growth association with climate was analysed at regional and local scales using correlation coefficient and bootstrapped response functions.

? Radial growth at both scales was positively correlated with rainfall during and prior to the growing season at all sites, and with summer rainfall before the growing season at five sites. Mean temperature effect changed according to the sampling site, from non-significant at the highest sites to significant (positive relationship in winter) at the lowest sites. Growth season temperature also had a negative effect.

? The Kalman filter was applied to estimate changing association between growth and climate over-time. Results suggested a change in association, initiated in the 80s, from non-significant to significant (*p < 0.05) at six of the sampling sites.

? Pinus pinaster is an accurate species for analysing tree-growth association with climate and for studying plant behaviour under global change conditions.

     

Resin tapping in Pinus pinaster: effects on growth and response function to climate

Pinus pinaster is of great ecological and economic importance and has traditionally been subjected to intensive uses such as wood and resin extraction. In the last decade, dendrochronological methods are increasingly being used to analyze the effects of climatic factors on the growth of the maritime pine, although tapped trees were generally avoided because it was thought that their growth was affected by resin extraction. In Spain, however, it is hard to find a long-lived forest of P. pinaster that has not been subjected to tapping for resin. In the present paper, we performed dendrochronological analyses of this species from wood cores and cross sections taken from both resin-tapped trees and resin-untapped trees killed by a fire in 2008 in the central Iberian region. On the one hand, we reconstructed the history of forest management by means of analysis of resin scars in the cross sections of resin-tapped trees. This facet of dendrochronological dating had not heretofore been developed, and little is therefore known about it. We dated 46 scars, which indicate a history of intensive resin extraction in the 1920–1950 period. Moreover, we attempted to answer the question: Have the old resin extractions in P. pinaster precluded the use of their growth rings for dendrochronological and dendroclimatic studies? We found that resin extraction did not alter general short-wavelength variability, and we developed a local chronology with all synchronized series, and the response function with respect to climate is similar to other oldest P. pinaster forests studied in Spain. The information we have recorded can be of use for providing tools to land managers for interpreting forest dynamics in resin-tapped regions.     

Plastic response of four maritime pine (<Emphasis Type="Italic">Pinus pinaster</Emphasis> Aiton) families to controlled soil water deficit

Key message

Separating the internal (ontogenetic) and external (environmental) components of maritime pine development during controlled soil water deficit helps to highlight the plastic response. The adjusted measurements reveal significant differences between families for their plastic response for several physiology and growth traits.

Context

Soil water deficit is and will be a growing problem in some regions. Pinus pinaster Ait. is a species of commercial interest and is recognized as a drought-avoiding species. It is thus of interest to evaluate the adaptation potential of P. pinaster to soil water deficit.

Aims

This paper aims to estimate the plastic response to the variation of water availability at the family level (half-sibs).

Methods

Two-year-old P. pinaster cuttings from four families were submitted during 6 weeks to two contrasting watering regimes. The experiment started in April 2011 shortly after sprouting. The photosynthesis and stomatal conductance to water vapor were measured on 1-year-old needles. Intrinsic water-use efficiency was calculated as the ratio of photosynthesis to stomatal conductance. Radial growth, length of terminal shoot, and total height were also measured. The ontogenetic component of tree development was estimated on the well-watered trees for all the traits. Then, this development effect was eliminated from the data collected on the trees submitted to the soil water deficit in order to keep only the effect of this soil water deficit.

Results

After 6 weeks of reduced watering, the value of all adjusted traits decreased. An average plastic response to the variation of water availability was found to be significant and variable at the family level for the six adjusted variables.

Conclusion

These results suggest that there is genetic variation of phenotypic plasticity to drought in P. pinaster for several traits, including stomatal conductance, which appears to be a promising variable for future selection for resistance to drought.

     

Acclimation of loblolly pine (Pinus taeda) seedlings to high temperatures

To determine the extent to which loblolly pine seedlings (Pinus taeda L.) acclimate to high temperatures, seedlings from three provenances-southeastern Texas (mean annual temperature 20.3 degrees C), southwestern Arkansas (mean annual temperature 16.2 degrees C) and Chesapeake, Maryland (mean annual temperature 12.8 degrees C)-were grown at constant temperatures of 25, 30, 35 or 40 degrees C in growth chambers. After two months, only 14% of the seedlings in the 40 degrees C treatment survived, so the treatment was dropped from the experiment. Provenance and family differences were not significant for most measured variables. Total biomass was similar in the 25 and 30 degrees C treatments, and less in the 35 degrees C treatment. Foliage biomass was higher, and root biomass lower, in the 30 degrees C treatment compared with the 25 degrees C treatment. Net photosynthesis and dark respiration of all seedlings were measured at 25, 30 and 35 degrees C. Both net photosynthesis and dark respiration exhibited acclimation to the temperature at which the seedlings were grown. For each temperature treatment, the highest rate of net photosynthesis was measured at the growth temperature. Dark respiration rates increased with increasing measurement temperature, but the basal rate of respiration, measured at 25 degrees C, decreased from 0.617 &mgr;mol m(-2) s(-1) in the 25 degrees C treatment to 0.348 &mgr;mol m(-2) s(-1) in the 35 degrees C treatment, resulting in less carbon loss in the higher temperature treatments than if the seedlings had not acclimated to the growth conditions. Temperature acclimation, particularly of dark respiration, may explain why total biomass of seedlings grown at 30 degrees C was similar to that of seedlings grown at 25 degrees C.     

Dynamic response of stomata to changing irradiance in loblolly pine (Pinus taeda L.)

Dynamic changes in stomatal conductance and the rate of photosynthesis were measured as periods of shading (decrease in irradiance from 800 to 200 micro mol m(-2) s(-1)) between 5 and 60 min were imposed on needles of Pinus taeda L. trees under laboratory conditions. Shading induced a 39% decrease in stomatal conductance but the rate of change was slow. Average time constants (+/- standard error) were shorter for the decrease in stomatal conductance when shading was imposed for 30 min (14.8 +/- 1.3 min) than for the increase in stomatal conductance when shading was removed (25.5 +/- 3.4 min). The time constants for increasing stomatal conductance when shading was removed were linearly related to the length of the previous dark period. The rate of photosynthesis fell immediately by 58% when shading was imposed and increased more rapidly than the change in stomatal conductance when shading was removed. The increase in photosynthesis during the induction phase after shading was removed was limited by both stomatal and biochemical effects. The long time constants for stomatal response contributed to the poor correlations between stomatal conductance and instantaneous measurements of irradiance from field data. However, the slow response of stomatal conductance to changes in irradiance had little effect on total daily transpiration, carbon gain and water-use efficiency.     

Genetic variation in susceptibility to pine wilt disease of maritime pine (<Emphasis Type="Italic">Pinus pinaster</Emphasis> Aiton) half-sib families

Key message

This paper presents a greenhouse study for assessing the genetic variation in maritime pine (Pinus pinaster Aiton) in response to pinewood nematode (Bursaphelenchus xylophilus (Steiner et Buhren) Nickle), which is a causal agent of pine wilt disease. Fifteen out of 96 half-sib families were selected as less susceptible. This experiment is an important first step for creating a resistance breeding program.

Context

Pine wilt disease is caused by the pinewood nematode (Bursaphelenchus xylophilus (Steiner et Buhren) Nickle), a quarantine pest, and is a concern to maritime pine (Pinus pinaster) in Portugal due to its economic, environmental, and social impacts. This disease is regarded as a major threat to European forests.

Aims

This paper aimed to evaluate the genetic variation in maritime pine families that were inoculated with pinewood nematode, identify the most resistant families, and establish the guidelines for a resistance improvement program.

Methods

Two-year-old half-sib progenies obtained from 96 plus trees were inoculated. The plants were monitored for survival on four different dates. The statistical analysis followed the mixed model theory.

Results

Genetic variability of the susceptibility to pine wilt disease was observed. At 157 days after inoculation, the 15 highest genetic ranking families out of 96 were selected, having a predicted survival mean of 15.6% instead of 11.0% on average for the all 96 families.

Conclusion

This study allows for the implementation of an improvement program to help control pine wilt disease.