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1.
? Premise of study: Prior work using a large data set has shown that the mechanical properties of wood disproportionately increase with increasing wood density across diverse species, e.g., stems composed of denser wood are stiffer and stronger than stems with equivalent cross-sections composed of less dense wood. However, an alternative approach, introducing the precondition of constant construction cost for the same data set, adduces that for any given construction cost, stems composed of lesser dense woods are stiffer and stronger then stems composed of denser woods. ? Methods: We evaluated these two approaches using generic allometric principles and the same large data set. ? Key results: This evaluation shows that construction costs cannot be constant over an entire ensemble of stems composed of different species of wood. For any specified construction cost (denoted by a k-value), only a particular subgroup of stems is addressed. The conclusions derived for this subgroup cannot be generalized to the entire ensemble of stems composed of different species of wood. ? Conclusion: Stems composed of denser wood are, on average as stiff and strong, or stiffer and stronger than stems with equivalent cross-sections composed of less dense wood. Denser wood may have a higher carbon construction cost, but its mechanical benefits likely outweigh the extra cost.  相似文献   

2.
? Premise of the study: In a previous paper, we questioned the traditional interpretation of the advantages and disadvantages of high wood density (Functional Ecology 24: 701-705). Niklas and Spatz (American Journal of Botany 97: 1587-1594) challenged the biomechanical relevance of studying properties of dry wood, including dry wood density, and stated that we erred in our claims regarding scaling. ? Methods: We first present the full derivation of our previous claims regarding scaling. We then examine how the fresh modulus of rupture and the elastic modulus scale with dry wood density and compare these scaling relationships with those for dry mechanical properties, using almost exactly the same data set analyzed by Niklas and Spatz. ? Key results: The derivation shows that given our assumptions that the modulus of rupture and elastic modulus are both proportional to wood density, the resistance to bending is inversely proportional to wood density and strength is inversely proportional with the square root of wood density, exactly as we previously claimed. The analyses show that the elastic modulus of fresh wood scales proportionally with wood density (exponent 1.05, 95% CI 0.90-1.11) but that the modulus of rupture of fresh wood does not, scaling instead with the 1.25 power of wood density (CI 1.18-1.31). ? Conclusions: The deviation from proportional scaling for modulus of rupture is so small that our central conclusion remains correct: for a given construction cost, trees with lower wood density have higher strength and higher resistance to bending.  相似文献   

3.
Wood density plays a central role in the life-history variation of trees, and has important consequences for mechanical properties of wood, stem and branches, and tree architecture. Wood density, modulus of rupture, modulus of elasticity, and safety factors for buckling and bending were determined for saplings of 30 Bolivian rain forest tree species, and related to two important life-history axes: juvenile light demand and maximum adult stature. Wood density was strongly positively related to wood strength and stiffness. Species safety factor for buckling was positively related to wood density and stiffness, but tree architecture (height : diameter ratio) was the strongest determinant of mechanical safety. Shade-tolerant species had dense and tough wood to enhance survival in the understorey, whereas pioneer species had low-density wood and low safety margins to enhance growth in gaps. Pioneer and shade-tolerant species showed opposite relationships between species traits and adult stature. Light demand and adult stature affect wood properties, tree architecture and plant performance in different ways, contributing to the coexistence of rain forest species.  相似文献   

4.
The ability of stem bark to resist bending forces was examined by testing in bending segments of Acer saccharum, Fraxinus americana, and Quercus robur branches with and without their bark. For each species, the bark contributed significantly to the ability of stem segments differing in age to resist bending forces, but its contribution was age-dependent and differed among the three species. The importance of the mechanical role of the bark decreased basipetally with increasing age of F. americana and Q. robur stem segments and was superceded by that of the wood for segments ≥ 6 yr old. A. saccharum bark was as mechanically important as the wood for stem segments 7 yr old but was not a significant stiffening agent for younger or older portions of stems. On average, the stiffness of the bark from all three species was 50% that of the wood. However, the geometric contribution to the flexural rigidity of stems made by the bark (i.e., the bark's second moment of area) was sufficiently large to offset its lower stiffness (Young's modulus) relative to that of the wood. A simple model is presented that shows that the bark must be as mechanically important as the wood when its radial thickness equals 32% that of the wood and its stiffness is 50% that of the wood. Based on this model, which is shown to comply with the data from three species purported to have stiff woods, it is evident that the role of the bark cannot be neglected when considering the mechanical behavior of juvenile woody stems subjected to externally applied bending forces.  相似文献   

5.
The allometric relationship of stem length L with respect to mean stem diameter D was determined for 80 shoots of each of three columnar cactus species (Stenocereus thurberi, Lophocereus schottii, and S. gummosus) to determine whether this relationship accords with that predicted by each of three contending models purporting to describe the mechanical architecture of vertical shoots (i.e., geometric, stress, and elastic similitude, which predict L proportional to D(alpha), with alpha = 1/1, 1/2, and 2/3, respectively). In addition, anatomical, physical, and biomechanical stem properties were measured to determine how the stems of these three species maintain their elastic stability as they increase in size. Reduced major axis regression of L with respect to D showed that alpha = 2.82 ± 0.14 for S. thurberi, 2.32 ± 0.19 for L. schottii, and 4.21 ± 0.31 for S. gummosus. Thus, the scaling exponents for the allometry of L differed significantly from that predicted by each of the three biomechanical models. In contrast, these exponents were similar to that for the allometry previously reported for saguaro. Analyses of biomechanical data derived from bending tests performed on 30 stems selected from each of the three species indicated that the bulk stem tissue stiffness was roughly proportional to L2, while stem flexural rigidity (i.e., the ability to resist a bending force) scaled roughly as L3. Stem length was significantly and positively correlated with the volume fraction of wood, while regression analysis of the pooled data from the three species (i.e., 90 stems) indicated that bulk tissue stiffness scaled roughly as the 5/3-power of the volume fraction of wood in stems. These data were interpreted to indicate that wood served as the major stiffening agent in stems and that this tissue accumulates at a sufficient rate to afford unusually high scaling exponents tot stem length with respect to stem diameter (i.e., disproportionately large increments of stem length with respect to increments in stem diameter). Nevertheless, the safety factor against the elastic failure of stems (computed on the basis of the critical buckling height divided by actual stem length) decreased with increasing stem size tot each species, even though each species maintained an average safety factor equal to two. We speculate that the apparent upper limit to plant height calculated for each species may serve as a biomechanical mechanism for vegetative propagation and the establishment of dense plant colonies by means of extreme stem flexure and ultimate breakage, especially for S. gummosus.  相似文献   

6.
The stems of large trees serve in transport, storage, and support; however, the degree to which these roles are reflected in their morphology is not always apparent. The large, water-filled stems of baobab trees (Adansonia spp.) are generally assumed to serve a water storage function, yet recent studies indicate limited use of stored water. Through an analysis of wood structure and composition, we examined whether baobab morphology reflects biomechanical constraints rather than water storage capacity in the six Madagascar baobab species. Baobab wood has a high water content (up to 79%), low wood density (0.09-0.17 g · cm(-3)), high parenchyma content (69-88%), and living cells beyond 35 cm into the xylem from the cambium. Volumetric construction cost of the wood is several times lower than in more typical trees, and the elastic modulus approaches that of parenchyma tissue. Safety factors calculated from estimated elastic buckling heights were low, indicating that baobabs are not more overbuilt than other temperate and tropical trees, yet the energy investment in stem material is comparable to that in temperate deciduous trees. Furthermore, the elastic modulus of the wood decreases with water content, such that excessive water withdrawal from the stem could affect mechanical stability.  相似文献   

7.
《Annals of botany》1997,79(5):479-485
The purpose of this paper is to determine the extent to which the physical and mechanical properties of dry and green wood samples are correlated. Samples of green (fresh) sap- and heartwood differing in density (ρ) were removed from the trunk of a black locust (Robinia pseudoacaciaL.) tree 30 years old and measuring 15 m in height. These samples were mechanically tested to determine their Young's elastic modulus (E), proportional (elastic) limit (σp), and modulus of rupture (σR). The Young's elastic modulus of green wood samples increased in magnitude to a limit with increasing cross-sectional area of the sample tested. The values of all three mechanical parameters measured for sapwood samples were consistently lower than those measured for heartwood samples with equivalent cross-sectional areas.Ewas linearly and positively correlated with the σpand σRof heartwood tissue samples. All mechanical properties were highly correlated with the density of green heartwood. Likewise, these properties were highly correlated with the specific gravity of wood samples. Based on these results, it is concluded that either the density of fresh wood or the specific gravity of air-dried wood can be used to estimate the mechanical properties of black locust wood based on simple regression curves in the absence of extensive mechanical tests.  相似文献   

8.
Obtaining representative values of green wood properties is essential for studies investigating the biomechanical aspects of tree development and ecology. Here, we compare the biomechanical properties of wood stored in various conditions between their collection in the field and their measurement. The study was performed on a large sample of wood specimens from different tropical species and different location in the trees, representing a wide diversity in wood structures. Elastic and viscoelastic properties are measured on green wood, and measured again after storage in different conditions: immersion in cold water during various durations, storage in an ethanol solution with or without washing in water, and air drying with or without rehydration. The systematic and random errors induced by these storage methods are quantified. Storage in cold water is the best way to preserve wood native properties. Soaking in ethanol is a fair alternative regarding elastic properties, but induces a significant change in viscoelastic properties. Air drying causes important, and partly irreversible, changes in mechanical properties. However, regarding elastic properties, this change is a systematic bias so that the air-dried elastic modulus provides a good basis for comparative studies of green wood stiffness.  相似文献   

9.
Aims Plant functional traits determine how plants respond to environmental factors and influence ecosystem processes. Among them, root traits and analyses of relations between above and below-ground traits in natural communities are scarce. Methods we characterized a set of above- and below-ground traits of three dominant shrub species in a semiarid shrub-steppe that had contrasting leaf phenological habits (deciduous, semideciduous and evergreen). We analysed if there was coordination among above- and below-ground resource economics patterns: i.e. patterns of biomass allocation, construction costs and lifespan.Important findings Above- and below-ground traits and their resource economics relations pointed to species-specific functional strategies to cope with drought and poor soils and to a species ranking of fast to slow whole-plant strategies in terms of resource uptake, biomass construction costs and turnover. The deciduous shrub, Proustia cuneifolia, had relatively deep and even distribution of roots, and high proportion of short-lived tissues of low C construction costs: it had high fine to coarse root and high leaf-to-stem biomass ratios, high specific leaf area (SLA), and stems of low wood density. This strategy allows Proustia to maximize and coordinate above- and below-ground resources uptake as long as the most limiting factor (water) is available, but at the cost of having relative high plant biomass turnover. The evergreen Porlieria chilensis, instead, displayed a more conservative and slow strategy in terms of resource economics. It had ~80% of the roots in the 40cm topsoil profile, low proportion of fine compared with coarse roots and low leaf-to-stem ratios, low SLA and stems of high wood density, i.e. it invested in C costly tissues that, overall, persist longer but probably at the cost of having lower plant resource uptake rates. Traits in the semideciduous Adesmia bedwellii were in between these two functional extremes. Our results revealed high functional diversity and above- and below-ground complementarity in resource economics among these three codominant species in the Chilean coastal desert.  相似文献   

10.
Tree architecture has important consequences for tree performance as it determines resource capture, mechanical stability and dominance over competitors. We analyzed architectural relationships between stem and crown dimensions for 13 dominant Iberian canopy tree species belonging to the Pinaceae (six Pinus species) and Fagaceae (six Quercus species and Fagus sylvatica) and related these architectural traits to wood density, shade tolerance and climatic factors. Fagaceae had, compared with Pinaceae, denser wood, saplings with wider crowns and adults with larger maximal crown size but smaller maximal height. In combination, these traits enhance light acquisition and persistence in shaded environments; thus, contributing to their shade tolerance. Pinaceae species, in contrast, had low-density wood, allocate more resources to the formation of the central trunk rather than to branches and attained taller maximal heights, allowing them to grow rapidly in height and compete for light following disturbances; thus, contributing to their high light requirements. Wood density had a strong relationship with tree architecture, with dense-wooded species having smaller maximum height and wider crowns, probably because of cheaper expansion costs for producing biomechanically stable branches. Species from arid environments had shorter stems and shallower crowns for a given stem diameter, probably to reduce hydraulic path length and assure water transport. Wood density is an important correlate of variation in tree architecture between species and the two dominant families, with potentially large implications for their resource foraging strategies and successional dynamics.  相似文献   

11.
? Wind is a key mechanical stress for woody plants, so how do shoot traits affect performance in wind? ? We used a vehicle mounted apparatus to measure drag, streamlining and mechanical safety in 127 vertical lead-shoots, 1.2 m long, across 39 species in tropical Australia. ? Shoot dimensions and stem tissue properties were closely coupled so that shoots with low stem specific gravity or larger projected area had thicker stems. Thicker stems provide larger second moment of area (I), which increased shoot safety and bending stiffness but impeded shoot reconfiguration in strong winds, including frontal area reduction. Nonetheless, increasing I also improved streamlining. Streamlining was unrelated to traits except I. Stem tissue material properties only had small effects. Higher modulus of rupture increased shoot safety and higher Young's modulus impeded shoot reconfiguration. ? We found no conflict between bending stiffness and streamlining for woody shoots. Stiffness might help streamlining by increasing damping and stability, thereby reducing flagging in wind. Tissue-level traits did influence shoot-level mechanical safety and behaviour, but shoot geometry was much more important. Variable shoot and stem traits, which all influenced shoot biomechanics, were integrated in shoots to yield a relatively narrow range of outcomes in wind.  相似文献   

12.
Violins produced by Antonio Stradivari during the late 17th and early 18th centuries are reputed to have superior tonal qualities. Dendrochronological studies show that Stradivari used Norway spruce that had grown mostly during the Maunder Minimum, a period of reduced solar activity when relatively low temperatures caused trees to lay down wood with narrow annual rings, resulting in a high modulus of elasticity and low density. The main objective was to determine whether wood can be processed using selected decay fungi so that it becomes acoustically similar to the wood of trees that have grown in a cold climate (i.e. reduced density and unchanged modulus of elasticity). This was investigated by incubating resonance wood specimens of Norway spruce (Picea abies) and sycamore (Acer pseudoplatanus) with fungal species that can reduce wood density, but lack the ability to degrade the compound middle lamellae, at least in the earlier stages of decay. Microscopic assessment of the incubated specimens and measurement of five physical properties (density, modulus of elasticity, speed of sound, radiation ratio, and the damping factor) using resonance frequency revealed that in the wood of both species there was a reduction in density, accompanied by relatively little change in the speed of sound. Thus, radiation ratio was increased from 'poor' to 'good', on a par with 'superior' resonance wood grown in a cold climate.  相似文献   

13.
Cuticle thickness of leaves varies >?100 times across species, yet its dry mass cost and ecological benefits are poorly understood. It has been repeatedly demonstrated that thicker cuticle is not superior as a water barrier, implying that other functions must be important. Here, we measured the mechanical properties, dry mass and density of isolated cuticle from 13 evergreen woody species of Australian forests. Summed adaxial and abaxial cuticle membrane mass per unit leaf area (CMA) varied from 2.95 to 27.4?g m(-2) across species, and accounted for 6.7-24% of lamina dry mass. Density of cuticle varied only from 1.04 to 1.24?g?cm(-3) ; thus variation in CMA was mostly due to variation in cuticle thickness. Thicker cuticle was more resistant to tearing. Tensile strength and modulus of elasticity of cuticle were much higher than those of leaf laminas, with significant differences between adaxial and abaxial cuticles. While cuticle membranes were thin, they could account for a significant fraction of leaf dry mass due to their high density. The substantial cost of thicker cuticle is probably offset by increased mechanical resistance which might confer longer leaf lifespans among evergreen species.  相似文献   

14.
Woody stems comprise a large biological carbon fraction and determine water transport between roots and leaves; their structure and function can influence both carbon and hydrological cycles. While angiosperm wood anatomy and density determine hydraulic conductivity and mechanical strength, little is known about interrelations across many species. We compiled a global data set comprising two anatomical traits for 3005 woody angiosperms: mean vessel lumen area (ā) and number per unit area (N). From these, we calculated vessel lumen fraction (F = āN) and size to number ratio (S = ā/N), a new vessel composition index. We examined the extent to which F and S influenced potential sapwood specific stem conductivity (K(S)) and wood density (D; dry mass/fresh volume). F and S varied essentially independently across angiosperms. Variation in K(S) was driven primarily by S, and variation in D was virtually unrelated to F and S. Tissue density outside vessel lumens (D(N)) must predominantly influence D. High S should confer faster K(S) but incur greater freeze-thaw embolism risk. F should also affect K(S), and both F and D(N) should influence mechanical strength, capacitance, and construction costs. Improved theory and quantification are needed to better understand ecological costs and benefits of these three distinct dimensions.  相似文献   

15.
The construction cost of a plant tissue, i.e. the amount of photoassimilates used in the synthesis of a unit weight, varies with its biochemical composition. Crop modellers use standard values published for a few groups of cultivated species. Yet, there are also intraspecific variations in the construction cost in relation with the development of the plant or organ. This research aimed at analysing the ontogenic changes in the construction cost of leaves, stems, roots, and fruits of tomato plants and the specific contribution of the mineral content to these changes. For that purpose, samples were harvested from the vegetative phase to the beginning of fruit production. The estimation of the construction cost was based on the contents of carbon, nitrogen and ash. In leaves, the construction cost decreased with the physiological age whereas, in stem internodes, it varied with the sympod number. These ontogenic changes could partly be explained by different accumulations of minerals. In contrast, the construction cost and the mineral content of fruits and roots remained fairly stable. On a whole plant basis, the construction cost of the bulk of each category of organs varied much less. Most of the increase in the mean construction cost of the whole plant during the experiment was due to changes in the allocation ratio between the vegetative parts and the fruits. Attention of crop modellers is drawn to the importance of a precise estimation of the construction cost and to the existence of ontogenic changes at the whole plant and organ levels.Key words: Lycopersicon esculentum Mill., construction cost, mineral content, ontogeny, carbon content   相似文献   

16.
The allometry of plant height H with respect to mean stem diameter D was determined based on 118 saguaro plants. The slope obtained for the reduced major axis regression analysis of the data was 2.36 ± 0.085, indicating that taller plants are disproportionately more slender than their shorter, presumably younger counterparts. The consequences of this positive, extremely anisometric relation on the elastic stability of stems were estimated by computing the critical buckling height Hcrit for each of the 118 stems on the basis of the mean density-specific stiffness (i.e., the quotient of Young's elastic modulus E and bulk tissue density ρ) determined for a single section from a mature saguaro stem. E/ρ was nearly equivalent to that of tissue samples of sclerenchyma isolated from other plant species. Since the slope of Hcrit vs. D equals ≈ 0.67 when E/ρ ≈ a constant, the safety-factor for saguaro stems (i.e., Hcrit/H) appeared to be size-dependent such that it decreased with increasing plant height (i.e., Hcrit/H ≈ D-1.65). However, the mean safety-factor computed for the 118 saguaro specimens was 9.64, indicating that, on the average, plant height was well below Hcrit. Additionally, circumstantial evidence suggests that saguaro stems become more stiff as they increase in size (and age) and that the rate of stem growth decelerates over time. The former would obtain a near size-independent safety-factor against elastic buckling while the latter protracts the time required to reach the critical buckling height. Comparisons among the allometries of H and Hcrit for saguaro, other cacti, nonwoody, and highly branched tree species indicated that saguaro size overlaps with the lower size-range of the largest known dicot and gymnosperm tree specimens likely as a consequence of the high E/ρ of mature saguaro stems.  相似文献   

17.
The physical and mechanical properties of wood affect the growth and development of trees, and also act as the main criteria when determining wood usage. Our understanding on patterns and controls of wood physical and mechanical properties could provide benefits for forestry management and bases for wood application and forest tree breeding. However, current studies on wood properties mainly focus on wood density and ignore other wood physical properties. In this study, we established a comprehensive database of wood physical properties across major tree species in China. Based on this database, we explored spatial patterns and driving factors of wood properties across major tree species in China. Our results showed that(i) compared with wood density, air-dried density, tangential shrinkage coefficient and resilience provide more accuracy and higher explanation power when used as the evaluation index of wood physical properties.(ii) Among life form, climatic and edaphic variables, life form is the dominant factor shaping spatial patterns of wood physical properties, climatic factors the next, and edaphic factors have the least effects, suggesting that the effects of climatic factors on spatial variations of wood properties are indirectly induced by their effects on species distribution.  相似文献   

18.
In eucalypt plantations managed for solid-wood products, radial trends in wood density, microfibril angle (MFA) and stiffness (modulus of elasticity, MoE) are properties of potential commercial importance that can be affected by competition from neighbouring trees. In this study, wood properties at breast height (1.3?m) were studied on radial strips prepared from 12-mm pith-to-bark wood cores taken from 20 trees in a 22-year-old Eucalyptus nitens (Deane and Maiden) Maiden thinning trial in north-eastern Tasmania, Australia. Thinning treatments were applied at age 6?years. Trees were sampled from each of the 200, 400?stems ha?1 and unthinned control treatments. Intra-specific competition for each sampled tree was estimated using the basal area growth of surrounding trees. SilviScan? technology was used to produce radial profiles of wood density, MFA and MoE. Results indicate a reduction in intra-specific competition through thinning of E. nitens plantations at an early age leads to a transient increase in MFA but has no significant effect on wood density or the intra-annual cycle of wood density. The correlation between the level of intra-specific competition and initial change in MFA following thinning, and a significant relationship between tree shape and mean MFA at breast height suggests the change in MFA is a post-thinning response to increased exposure and wind sway.  相似文献   

19.
Aims A better understanding of wood litter decomposition is essential for predicting responses of forest ecosystems to global climate change. Recent studies suggest that chemical properties of wood litters, rather than physical ones such as wood density, are more important for interspecific differences in wood decomposition rates. However, empirical data are still limited, especially for tropical trees. In addition, decomposition rate of wood litter often varies with time, which makes interspecific comparison difficult. We studied the wood decomposition of 32 rainforest trees to elucidate (i) the degree of interspecific variation in wood decomposition rate of a given size and configuration and (ii) if initial wood density and water permeability are consistent predictors of the overall decomposition rate and its pattern over time.Methods A common garden decomposition experiment was conducted in a tropical rainforest in Malaysian Borneo for 32 native tree species. Small wood sticks were set on the forest floor and the weight loss was monitored monthly for 2.7 years.Important findings We found large variation in the wood decomposition rate (a 49-fold range), suggesting that we need to consider this variation when calculating community-level carbon dynamics of tropical rain forests. The physical traits of wood, i.e. wood density and water permeability, were related to wood decomposition rate and its pattern over time. Decomposition half-time related positively and negatively to initial wood density and water permeability, respectively. The time-dependent-rate model fitted better for 18 species (56% of the study species) that had higher water permeabilities than the others, suggesting that micelle porosity in wood relates to temporal changes in decomposition rate.  相似文献   

20.
Aim Wood properties are related to tree physiology and mechanical stability and are influenced by both phylogeny and the environment. However, it remains unclear to what extent geographical gradients in wood traits are shaped by either phylogeny or the environment. Here we aimed to disentangle the influences of phylogeny and the environment on spatial trends in wood traits. Location China. Methods We compiled a data set of 11 wood properties for 618 tree species from 98 sampling sites in China to assess their phylogenetic and spatial patterns, and to determine how many of the spatial patterns in wood properties are attributable to the environment after correction for phylogenetic influences. Result All wood traits examined exhibited significant phylogenetic signal. The widest divergence in wood traits was observed between gymnosperms and angiosperms, Rosids and Asterids, Magnoiliids and Eudicots, and in Lamiales. For most wood traits, the majority of trait variation was observed at genus and species levels. The mechanical properties of wood showed correlated evolution with wood density. Most of the mechanical properties of wood exhibited significant latitudinal variation but limited or no altitudinal variation, and were positively correlated with mean annual precipitation based on both Pearson's correlation analysis and the phylogenetic comparative method. Correlations at family level between mean annual temperature and wood density, compression strength, cross‐section hardness, modulus of elasticity and volumetric shrinkage coefficient became significant after correction for phylogenetic influences. Main conclusions Phylogeny interacted with the environment in shaping the spatial patterns of wood traits of trees across China because most wood properties showed strong phylogenetic conservatism and thus affected environmental tolerances and distributions of tree species. Mean annual precipitation was a key environmental factor explaining the spatial patterns of wood traits. Our study provides valuable insights into the geographical patterns in productivity, distribution and ecological strategy of trees linking to wood traits.  相似文献   

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