Adaptive anatomy of Pinus halepensis trees from different Mediterranean environments in Spain

A study was conducted on the variation in growth, biomass, juvenile wood anatomy, and needle morphology of Pinus halepensis Mill. from three Spanish regions of provenance characterized by environmental differences, without the influence of the site factor. Seeds collected from two progenies in each region were planted in a single plot, and the trees were felled at the age of 7 years. The results showed significant differences between provenances, as well as the genetic or environmental influence on the traits analyzed. Trees adapted to moderate summer drought conditions (Inland Catalonia region) are primarily characterized by higher average values for height, diameter, biomass, cell wall thickness, inter-tracheid wall strength, pit membrane diameter, torus diameter, bordered pit aperture diameter, and ray tracheid abundance in comparison with trees adapted to severe summer drought conditions (Southern region and Balearic Islands region). The greater structural requirements of trees from the Inland Catalonia region, subjected to higher weight and wind loads, resulted in thick cell walls. Moreover, the large pits and more abundant ray tracheids in trees from this provenance would allow more efficient water transport and greater water storage capacity, respectively. The differences found between provenances suggest the adaptive nature of the anatomy of this species, which demonstrates the importance of the region of provenance when choosing reproduction material for reforestation.     

红松种源材质性状研究

对帽儿山地区17 年生26 个红松种源试验林进行了材质性状研究, 主要包括材质性状在不同种源间遗传变异情况以及从髓心到树皮径向变异模式研究。结果表明:红松各项材质在不同种源间均存在着极显著差异。各项材质均存在着一定的径向变异模式, 主要表现在:红松各种源的管胞长度、管胞直径、木材密度和晚材率总体趋势是从髓心向外呈迅速增加趋势, 而年轮宽从髓心向外呈递减趋势。本项研究为红松材性早期选择和预测提供依据。     

Intra-annual tracheid formation of Norway spruce provenances in southern Finland

We studied the intra-annual wood formation in a Norway spruce provenance experiment in southern Finland from 2004–2008. Two Finnish provenances, northern and southern, as well as German and Hungarian provenances were included. Timing of tracheid formation and differentiation, and tracheid dimensions were determined from periodically extracted microcores. The aim was to determine the differences between the years and provenances in the timing of the xylogenesis and in the xylem characteristics. Year-to-year variation was high both in timing of tracheid formation and xylem characteristics, while between-provenance differences were small. The onset of tracheid formation varied from early May to late June in different trees in different years. The onset of tracheid formation was not closely related to the annual variations of temperature sum. In all the years, daily temperatures exceeded the threshold +5°C for several weeks before the onset of tracheid formation. The highest tracheid formation rate occurred after the summer solstice in all years and generally coincided with the highest daily temperatures during the growing season. Tracheid production ceased early in 2006 due to a mid-summer drought. Cell differentiation continued late in autumn as non-mature tracheids were still observed around mid-September. No clear differences between the provenances in the timing of tracheid formation were observed, although the Finnish provenances tended to initiate tracheid formation slightly earlier than the other provenances. The tree-ring widths of the Finnish provenances were also wider, while tracheid diameter of the German provenance was slightly smaller. Our results indicate that between-tree variation in the timing of wood formation is high compared with the latitude effect of seed source.     

Plastic Response of Tracheids in Pinus pinaster in a Water-Limited Environment: Adjusting Lumen Size instead of Wall Thickness

The formation of wood results from cambial activity and its anatomical properties reflect the variability of environmental conditions during the growing season. Recently, it was found that wood density variations in conifers growing under cold-limited environment result from the adjustment of cell wall thickness (CWT) to temperature. Additionally, it is known that intra-annual density fluctuations (IADFs) are formed in response to precipitation after the summer drought. Although IADFs are frequent in Mediterranean conifers no study has yet been conducted to determine if these structures result from the adjustment of lumen diameter (LD) or CWT to soil water availability. Our main objective is to investigate the intra-ring variation of wood anatomical features (LD and CWT) in Pinus pinaster Ait. growing under a water-limited environment. We compared the tracheidograms of LD and CWT for the years 2010–2013 in P. pinaster growing in the west coast of Portugal. Our results suggest a close association between LD and soil moisture content along the growing season, reinforcing the role of water availability in determining tracheid size. Compared with CWT, LD showed a higher intra- and inter-annual variability suggesting its strong adjustment value to variations in water availability. The formation of a latewood IADF appears to be predisposed by higher rates of cell production in spring and triggered by early autumn precipitation. Our findings reinforce the crucial role of water availability on cambial activity and wood formation in Mediterranean conifers, and emphasize the high plasticity of wood anatomical features under Mediterranean climate.     

Intra-annual variability of anatomical structure and δ<Superscript>13</Superscript>C values within tree rings of spruce and pine in alpine,temperate and boreal Europe

Tree-ring width, wood density, anatomical structure and ¹³C/¹²C ratios expressed as δ¹³C-values of whole wood of Picea abies were investigated for trees growing in closed canopy forest stands. Samples were collected from the alpine Renon site in North Italy, the lowland Hainich site in Central Germany and the boreal Flakaliden site in North Sweden. In addition, Pinus cembra was studied at the alpine site and Pinus sylvestris at the boreal site. The density profiles of tree rings were measured using the DENDRO-2003 densitometer, δ¹³C was measured using high-resolution laser-ablation-combustion-gas chromatography-infra-red mass spectrometry and anatomical characteristics of tree rings (tracheid diameter, cell-wall thickness, cell-wall area and cell-lumen area) were measured using an image analyzer. Based on long-term statistics, climatic variables, such as temperature, precipitation, solar radiation and vapor pressure deficit, explained <20% of the variation in tree-ring width and wood density over consecutive years, while 29–58% of the variation in tree-ring width were explained by autocorrelation between tree rings. An intensive study of tree rings between 1999 and 2003 revealed that tree ring width and δ¹³C-values of whole wood were significantly correlated with length of the growing season, net radiation and vapor pressure deficit. The δ¹³C-values were not correlated with precipitation or temperature. A highly significant correlation was also found between δ¹³C of the early wood of one year and the late wood of the previous year, indicating a carry-over effect of the growing conditions of the previous season on current wood production. This latter effect may explain the high autocorrelation of long-term tree-ring statistics. The pattern, however, was complex, showing stepwise decreases as well as stepwise increases in the δ¹³C between late wood and early wood. The results are interpreted in the context of the biochemistry of wood formation and its linkage to storage products. It is clear that the relations between δ¹³C and tree-ring width and climate are multi-factorial in seasonal climates.     

Studies on Wood Anatomy of Eucalyptus globulus Under Different Habitats

Wood anatomy of Eucalyptus globulus Labill. under different habitats was comparatively studied under light and scanning electron microscope. According to the eleven major quantitative perimeters of wood characteristics of E. globulus viz: vessel frequency, percentage of solitary vessels, vessel diameter, vessel member length; tracheid length and diameter; fibre tracheid length and diameter; libriform fibre length; ray frequency and ray height, the plants under different habitats have been made a comparison. The results showed that growth ring was indistinct, and there was a positive correlation between the quantitative characters of most wood constituent members were positively correlated with the latitude except for some of vessel frequency decreased. The decrease in effects of annual rainfall on the structure of the wood of E. glotrulus were evaluated.     

不同生境下蓝桉的木材解剖研究

用光镜及扫描电镜对不同生境下的蓝桉（Ｅｕｃａｌｙｐｔｕｓｇｌｏｂｕｌｕｓ Ｌａｂｉｌｌ．）木材结构进行了解剖学观察,描述了木材结构特征。以蓝桉木材的１１ 个主要特征——导管频率、单孔率、导管直径、导管分子长度、管胞长度和直径、纤维管胞长度和直径、纤维长度、射线频率和高度为指标,对生长在不同生境下的蓝桉进行比较发现：随着纬度的增加,生长轮变化不明显;木材中大部分组成分子的数量特征,除导管频率有降低趋势外均与纬度成正相关。评述了年降雨量对蓝桉木材结构的影响     

长白落叶松半同胞家系生长和木材性状遗传变异与联合选择

长白落叶松（Larix olgensis）是我国东北重要的用材树种,根据生长和木材性状对其进行综合选择至关重要。本研究以吉林省延边自治区汪清林业局32年生的49个长白落叶松半同胞家系为材料,对其9个生长性状（树高、地径、胸径、3 m径、材积、尖削度、冠幅、分枝角和通直度）和4个木材性状（木材基本密度、管胞长度、管胞宽度和管胞长宽比）进行测定与分析。结果表明：不同变异来源间所有性状差异均达极显著水平（P<0.01）;各性状家系遗传力均较高（0.51~0.96）;表型变异系数为3.04%（分枝角）~23.15%（冠幅）;各性状相关系数为-0.367（管胞宽度与管胞长宽比）~0.994（胸径与材积）;主成分分析结果表明,4个主成分的累计贡献率达到78.46%,包含了家系生长性状和木材性状的大部分信息;分别以生长和木材性状对家系进行综合评价,初步筛选出5个生长性状优良家系（S78、S81、S80、S84和S83）和5个木材性状优良家系（S37、S51、S6、S30和S19）,结合生长和木材性状初步筛选出5个优良家系（S89、S74、S76、S82和S83）。本研究初选的材料可以为良种选育提供基础,亲本可以为改良种子园营建提供材料。     

Daily dynamics in xylem cell radial growth of Scots pine (Pinus sylvestris L.)

Daily dynamics of radial cell expansion during wood formation within the stems of 25-year-old Scots pine trees (Pinus sylvestris L.), growing in field conditions, were studied. The samples of forming wood layers were extracted 4 times per day for 3 days. Possible variations in the growth on different sides of the stem, duration of cell development in radial cell expansion phase and dynamics of cell growth in this phase were taken into account. The perimeters of tracheid cross-sections as a reflection of primary cell wall growth were the criterion of growth in a radial direction. For the evaluation of growing cell perimeters a special system for digital processing and image analysis of tracheid cross-sections of the forming wood was used. Growth rate for certain time intervals was estimated by the change in the relation of the perimeter of each observed cell in each of ten tracheid rows in each of 12 trees to the perimeter of the xylem cell of the same row before the expansion. Temporal differences in average values of the relations were estimated by Analyses of Variance. The existence of daily dynamics of Scots pine xylem cell radial growth has been proved. Intensive growth of pine tracheids has been shown to occur at any time of the day and to depend on the temperature regime of the day and the night as well as water supply of stem tissues. Moreover, reliable differences (P = 0.95) in the increment of cell walls during tracheid radial expansion have been found. Pulsing changes of the water potentials both of the cell and the apoplast, as the reason for the fluctuations of radial cell growth rate, were discussed.     

MULTIVARIATE COMPARISON OF FOSSIL AND RECENT GINKGO WOOD

Multivariate statistical analysis of the anatomical structure of the wood of Recent Ginkgo biloba L. and of fossil Ginkgo beckii Scott, Barghoorn, & Prakash revealed significant differences not apparent through general microscopic observation. On the basis of a multiple discriminant analysis by computer of 10 anatomical characters, the living and fossil Ginkgo woods were separated into two distinct groups. All Recent wood specimens were classified into a single disjoint group by the computer, and all fossil wood specimens were classified into another without misclassifications when all 10 characteristics were included in the analysis. Three characters—number of pits per millimeter of tracheid length on radial wall, ray height, and tangential diameter of late wood tracheids—accounted for most of the discrimination between the two groups.     

我国南部5种冷杉植物木材的比较解剖学研究

应用光学显微镜和扫描电子显微镜对分布于我国南部的5种冷杉属(Abies)植物,即资源冷杉(A．ziyuanensis)、大院冷杉(A.dayuanensis)、梵净山冷杉(A.fanjingshanensis)、元宝山冷杉(A.yuanbaoshanensis)和台湾冷杉(Akawakamii)的木材结构进行了研究。从管胞分子长度、管胞分子直径、射线高度、射线频率、纹孔类型、交叉场纹孔类型、交又场纹孔数目等方面,比较了它们木材解剖特征的差异。结果表明：大院冷杉和资源冷杉在射线频率、管胞直径、管胞长度、交叉场纹孔数目方面都很类似;元宝山冷杉在管胞直径与射线长度方面与其它冷杉有显著差异。在纹孔类型、交叉场纹孔类型方面,几种冷杉都是一致的。在交叉场纹孔数目方面,大院冷杉、资源冷杉较少,而梵净山冷杉、元宝山冷杉和台湾冷杉较多。我国南部这5种冷杉的木材解剖特征与分布在华北、东北及青藏高原上冷杉的木材非常类似。为了便于区别这5种木材,根据木材解剖特征编制了检索表。     

A new method for vulnerability analysis of small xylem areas reveals that compression wood of Norway spruce has lower hydraulic safety than opposite wood

Compression wood is formed at the underside of conifer twigs to keep branches at their equilibrium position. It differs from opposite wood anatomically and subsequently in its mechanical and hydraulic properties. The specific hydraulic conductivity (k_s) and vulnerability to drought‐induced embolism (loss of conductivity versus water potential ψ) in twigs of Norway spruce [Picea abies (L.) Karst.] were studied via cryo‐scanning electron microscope observations, dye experiments and a newly developed ‘Micro‐Sperry’ apparatus. This new technique enabled conductivity measurements in small xylem areas by insertion of syringe cannulas into twig samples. The hydraulic properties were related to anatomical parameters (tracheid diameter, wall thickness). Compression wood exhibited 79% lower k_s than opposite wood corresponding to smaller tracheid diameters. Vulnerability was higher in compression wood despite its narrower tracheids and thicker cell walls. The P₅₀ (ψ at 50% loss of conductivity) was ?3.6 MPa in opposite but only ?3.2 MPa in compression wood. Low hydraulic efficiency and low hydraulic safety indicate that compression wood has primarily a mechanical function.     

Ecological wood anatomy of NepaleseRhododendron (Ericaceae). 1. Interspecific variation

The relationship of selected wood anatomical characters of NepaleseRhododendron with stem diameter, plant height, altitude, and plant form was investigated. We studied one to three specimens each of 26 species: five species each of trees and subtrees I, three species of subtrees II, and 13 species of shrubs. Multiple regression analysis and actual distribution of character values show that pore characters and multiseriate ray ratio have a stronger correlation with stem diameter than altitude; that pore density, vessel element length, fiber-tracheid length, and multiseriate ray density and width are equally related to altitude and stem diameter, or to altitude and plant height; and that bar number, and multiseriate ray area and height have a stronger connection with altitude. Among the characters, average pore area is most strongly correlated with stem diameter and increases exponentially as diameter increases. For wood structure of NepaleseRhododendron, 17 to 63 % of the variation is affected by non-anatomical factors. The general trends in wood structure of NepaleseRhododendron show that trees and subtrees form one continuous unit whereas shrubs form another that often has wider ranges of variation.     

Drought-induced increase in water-use efficiency reduces secondary tree growth and tracheid wall thickness in a Mediterranean conifer

In order to understand the impact of drought and intrinsic water-use efficiency (iWUE) on tree growth, we evaluated the relative importance of direct and indirect effects of water availability on secondary growth and xylem anatomy of Juniperus thurifera, a Mediterranean anisohydric conifer. Dendrochronological techniques, quantitative xylem anatomy, and ¹³C/¹²C isotopic ratio were combined to develop standardized chronologies for iWUE, BAI (basal area increment), and anatomical variables on a 40-year-long annually resolved series for 20 trees. We tested the relationship between iWUE and secondary growth at short-term (annual) and long-term (decadal) temporal scales to evaluate whether gains in iWUE may lead to increases in secondary growth. We obtained a positive long-term correlation between iWUE and BAI, simultaneously with a negative short-term correlation between them. Furthermore, BAI and iWUE were correlated with anatomical traits related to carbon sink or storage (tracheid wall thickness and ray parenchyma amount), but no significant correlation with conductive traits (tracheid lumen) was found. Water availability during the growing season significantly modulated tree growth at the xylem level, where growth rates and wood anatomical traits were affected by June precipitation. Our results are consistent with a drought-induced limitation of tree growth response to rising CO₂, despite the trend of rising iWUE being maintained. We also remark the usefulness of exploring this relationship at different temporal scales to fully understand the actual links between iWUE and secondary growth dynamics.     

Radial stem variations of Tabebuia chrysantha (Bignoniaceae) in different tropical forest ecosystems of southern Ecuador

Stem diameter increments of the broadleaved deciduous tree species Tabebuia chrysantha were measured with high-resolution dendrometers in a tropical lower montane forest and in a dry forest in southern Ecuador, the latter showing a distinct dry season. Those analyses were complemented by wood anatomical studies on regularly collected microcores to determine the season of active cambial growth and the time of formation of annual growth boundaries. The length of the cambial active period varied between 3 and 7 months at the tropical lower montane forest and 2 and 4 months in the dry forest, respectively. During dry days, amplitudes of daily stem diameter variations correlated with vapour pressure deficit. During October and November, inter-annual climate variations may lead to dry and sunny conditions in the tropical lower montane forest, causing water deficit and stem diameter shrinkage in T. chrysantha. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall as well as vapour pressure deficit in certain periods of the year, indicating that rainfall plays a major role for tree growth.     

Seasonal gas exchange and water relations in juveniles of two evergreen Neotropical savanna tree species with contrasting regeneration strategies

Colonization dynamics of woody species into grasslands in Neotropical savannas are determined by two main factors: plant-available moisture and fire. Considering seasonality of precipitation and high fire frequency in these ecosystems, vegetative reproduction has been suggested as the main regeneration strategy in woody species. This study examined seasonal variations in water relations and photosynthesis in juveniles of two tree species with contrasting regeneration strategies: Palicourea rigida (sexual reproduction) and Casearia sylvestris (asexual reproduction). The studied species showed similar transpiration rates to deep-rooted adult evergreen tree species during the rainy period, suggesting little water availability limitations on surface soil layers. P. rigida juveniles significantly decreased their leaf water potentials from wet to dry seasons. In C. sylvestris resprouts, there were no seasonal differences in their predawn water potentials and gas exchange parameters, indicating a water deficit avoidance characteristic derived from their connections to deep-rooted adult counterparts allowing access to moist soil at depth even during the drought period. P. rigida rely on strict control of water losses and turgor maintenance through elastic cell walls during the dry season. The iso-hydric behavior of gas exchange and most water relations parameters in C. sylvestris enable turgor maintenance during the dry season which also gives the possibility to achieve foliar expansion under water-stressed conditions for shallow-rooted plants. Nevertheless, in absence of water deficits, P. rigida had the advantage to be physiologically independent individuals, showing an equal or even superior photosynthetic performance that eventually could be translated into a more favorable whole-plant carbon balance and higher growth rates in wet habitats.     

Direct observation and modelling of embolism spread between xylem conduits: a case study in Scots pine

Xylem embolism is one of the main processes involved in drought‐related plant mortality. Although its consequences for plant physiology are already well described, embolism formation and spread are poorly evaluated and modelled, especially for tracheid‐based species. The aim of this study was to assess the embolism formation and spread in Pinus sylvestris as a case study using X‐ray microtomography and hydraulics methods. We also evaluated the potential effects of cavitation fatigue on vulnerability to embolism and the micro‐morphology of the bordered pits using scanning electron microscopy (SEM) to test for possible links between xylem anatomy and embolism spread. Finally, a novel model was developed to simulate the spread of embolism in a 2D anisotropic cellular structure. Results showed a large variability in the formation and spread of embolism within a ring despite no differences being observed in intertracheid pit membrane anatomical traits. Simulations from the model showed a highly anisotropic tracheid‐to‐tracheid embolism spreading pattern, which confirms the major role of tracheid‐to‐tracheid air seeding to explain how embolism spreads in Scots pine. The results also showed that prior embolism removal from the samples reduced the resistance to embolism of the xylem and could result in overestimates of vulnerability to embolism.     

Hydraulic efficiency compromises compression strength perpendicular to the grain in Norway spruce trunkwood

The aim of this study was to investigate bending stiffness and compression strength perpendicular to the grain of Norway spruce (Picea abies (L.) Karst.) trunkwood with different anatomical and hydraulic properties. Hydraulically less safe mature sapwood had bigger hydraulic lumen diameters and higher specific hydraulic conductivities than hydraulically safer juvenile wood. Bending stiffness (MOE) was higher, whereas radial compression strength lower in mature than in juvenile wood. A density-based tradeoff between MOE and hydraulic efficiency was apparent in mature wood only. Across cambial age, bending stiffness did not compromise hydraulic efficiency due to variation in latewood percent and because of the structural demands of the tree top (e.g. high flexibility). Radial compression strength compromised, however, hydraulic efficiency because it was extremely dependent on the characteristics of the “weakest” wood part, the highly conductive earlywood. An increase in conduit wall reinforcement of earlywood tracheids would be too costly for the tree. Increasing radial compression strength by modification of microfibril angles or ray cell number could result in a decrease of MOE, which would negatively affect the trunk’s capability to support the crown. We propose that radial compression strength could be an easily assessable and highly predictive parameter for the resistance against implosion or vulnerability to cavitation across conifer species, which should be topic of further studies.     

Declining hydraulic performances and low carbon investments in tree rings predate Scots pine drought-induced mortality

Key message

The retrospective analysis of wood anatomical features evidences how a long-term deterioration of hydraulic performance and carbon use portend drought-induced mortality in Scots pine.

Abstract

Widespread episodes of drought-induced tree mortality are predicted to become more frequent as climate becomes warmer and drier. Nevertheless, growth trends and their links to changes in wood anatomy before tree dies are still poorly understood. Wood anatomical features provide valuable information that can be extracted to infer the mechanisms leading to tree death. In this study, we characterize drought-induced mortality affecting two Scots pine (Pinus sylvestris) sites (Prades and Arcalís) located in the North Eastern Iberian Peninsula. Co-occurring now-dead and living Scots pine trees were sampled and their wood anatomical features were measured and compared. We aimed to detect differences in anatomical features between living and dead trees, and to infer past physiological performances that might have determined their subsequent death or survival. Now-dead trees showed lower tracheid and resin duct production, and smaller radial lumen diameters than co-occurring living trees. At the more xeric Prades site, these anatomical differences were larger and chronic, i.e. were observed over the three studied decades, whilst they were less pronounced at the other, more mesic Arcalís site, where tree mortality episodes were more recent. This indicates that dead trees’ hydraulic conductivity was severely affected and that carbon investment in xylem formation and resin duct production was constrained prior to tree death. Our findings show that both hydraulic deterioration and low carbon allocation to xylem formation were associated to drought-induced mortality in Scots pine. Nevertheless, the temporal dynamics of these processes differed between populations as a function of site climatic conditions.