Clara Tinoco-Ojanguren

Wood biomechanics and anatomy of PACHYCEREUS PRINGLEI.

We report the longitudinal, biomechanical, and anatomical trends observed for tissue samples drawn from the parallel aligned, prismatic woody vascular bundles running the length of a Pachycereus pringlei plant measuring 5.22 m in height. The main vertical stem of this plant was cut into five segments (labeled A through E in the acropetal direction) measuring ∼1.02 m in length. Four of the 14 vascular bundles in each segment were surgically removed to obtain 20 vascular bundle segments that were tested in bending to determine their stiffness measured in the radial E(R) and tangential E(T) direction. We also determined the lignin content of representative samples of wood.A nonlinear trend in stiffness was observed: E(R) and E(T) were highest in segments B or C (1.67 GN/m and 1.09 GN/m, respectively), lower in segment A (E(R) = 1.18 GN/m and E(T) = 0.35 GN/m), and lowest in segment E (E(R) = 0.03 GN/m and E(T) = 0.20 GN/m). Similar longitudinal trends were seen for axial tissue volume fraction and fiber wall thickness, which achieved their highest values in segment B (69.8% and 6.59 μm, respectively). Wood stiffness also correlated significantly with cell wall lignin content: with respect to segment B (which had the highest lignin content, and was thus used as the standard reference for percent lignin content), lignin content, was 15, 60, 85, and 43% in segments E, D, C, and A, respectively. Fiber cell length, which increased toward the base of the stem and toward the vascular cambium in the most proximal vascular bundle segment, did not correlate with E(R) or E(T).Basic engineering principles were used to calculate stem stresses resulting from self-loading and any wind-induced bending moment (produced by drag forces). Calculations indicated that the less stiff wood produced in segment A eliminates a rapid and potentially dangerous increase in stresses that would otherwise occur in segments B or C. The less stiff wood in segment A also reduces the probability of shear failure at the cellular interface between the wood and surrounding tissues in this portion of the stem.We conclude that P. pringlei wood stiffness is dependent on the volume fraction and lignification of axial tissues, less so on fiber wall thickness, and that wood development in this species is adaptively responsive to self-loading and differentially applied external mechanical forces.

Morphological and physiological differentiation of seedlings between dry and wet habitats in a tropical dry forest.

A common observation in tropical dry forests is the habitat preference of tree species along spatial soil water gradients. This pattern of habitat partitioning might be a result of species differentiation in their strategy for using water, along with competing functions such as maximizing water exploitation and tolerating soil water stress. We tested whether species from drier soil conditions exhibited a tolerance strategy compared with that of wet-habitat species. In a comparison of 12 morphophysiological traits in seedlings of 10 closely related dry and wet-habitat species pairs, we explored what trade-offs guide differentiation between habitats and species. Contrary to our expectations, dry-habitat species showed mostly traits associated with an exploitation strategy (higher carbon assimilation capacity, specific leaf area and leaf-specific conductivity and lower water-use efficiency). Strikingly, dry-habitat species tended to retain their leaves longer during drought. Additionally, we detected multiple strategies to live within each habitat, in part due to variation of strategies among lineages, as well as functional differentiation along the water storage capacity-stem density (xylem safety) trade-off. Our results suggest that fundamental trade-offs guide functional niche differentiation among tree species expressed both within and between soil water habitats in a tropical dry forest.

The biomechanics of Pachycereus pringlei root systems.

We report on the root system of the large columnar cactus species Pachycereus pringlei to explore the hypothesis that increasing plant size decreases the ability to resist wind-throw but increases the capacity to absorb and store nutrients in roots (i.e., plant size limits the performance of these functions and may shift the performance of one function in favor of another as size increases). Based on 18 plants differing in size, the root system is characterized by a broad and deep bayonet-like root central to a shallow and extensive lateral system of root elements bearing sinker roots near the stem base. All root types have a living secondary cortex and contain wood with a large volume fraction of ray tissues that increases toward the stem base. Wood stiffness and tensile strength are correlated negatively with the ray tissue volume fraction and thus decrease toward the stem base in lateral and bayonet roots. Calculations show that the ability of the bayonet and proximal lateral root elements to resist wind-throw decreases with increasing plant size, whereas the nutrient absorption/storage capacity of the total root system increases with plant size (i.e., a size-dependent shift between these two root functions occurs).

Biomechanics of the columnar cactus Pachycereus pringlei.

We report the longitudinal variations in stiffness and bulk density of tissue samples drawn from along the length of two Pachycereus pringlei plants measuring 3.69 and 5.9 m in height to determine how different tissues contribute to the mechanical stability of these massive vertical organs. Each of the two stems was cut into segments of uniform length and subsequently dissected to obtain and mechanically test portions of xylem strands, stem ribs, and a limited number of pith and cortex samples. In each case, morphometric measurements were taken to determine the geometric contribution each tissue likely made to the ability of whole stems to resist bending forces. The stiffness of each xylem strand increased basipetally toward the base of each plant where stiffness sharply decreased, reaching a magnitude comparable to that of strands 1 m beneath the stem apex. The xylem was anisotropic in behavior, i.e., its stiffness measured in the radial and in the tangential directions differed significantly. Despite the abrupt decrease in xylem strand stiffness at the stem base, the contribution made by this tissue to resist bending forces increased exponentially from the tip to the base of each plant due to the accumulation of wood. A basipetal increase in the stiffness of the pith (and, to limited extent, that of the cortex) was also observed. In contrast, the stiffness of stem rib tissues varied little as a function of stem length. These tissues were stiffer than the xylem in the corresponding portions of the stem along the upper two-fifths of the length of either plant. Tissue stiffness and bulk density were not significantly correlated within or across tissue types. However, a weak inverse relationship was observed for these properties in the case of the xylem and stem rib tissues. We present a simple formula that predicts when stem ribs rather than the xylem strands serve as the principal stiffening agents in stems. This formula successfully predicted the observed aspect ratio of the stem ribs (the average quotient of the radial and tangential dimensions of rib transections), and thus provided circumstantial evidence that the ribs are important for mechanical stability for the distal and younger regions of the stems examined.

Seasonal balance and vertical pattern of photosynthetically active radiation within canopies of a tropical dry deciduous forest ecosystem in Mexico

Major components of the flux density of global photosynthetically active radiation (PAR) were measured above and within canopies in a tropical deciduous forest on the Pacific coast of Mexico. At each of 69 locations grouped along a topographic sequence the PAR reflected from the top of the canopy, the vertical profile of transmittance, and the reflectance from the ground, were measured as many as four times in the year, including the extremes of the wet and dry seasons. With these observations an annual balance of the portion of PAR radiation reflected and absorbed by the canopy and ground was assembled and the detailed spatial and temporal dynamics of PAR within canopy layers were estimated. Canopy stature declined along the topographic sequence and the shape of the transmittance profiles reflected this. In locations of declining moisture availability the fraction of PAR absorbed by the ground increased and the fraction absorbed by non-foliar tissues decreased. Seasonal variation in canopy structure was the dominant influence on the partitioning of radiation - spatial variation was less important. Of a total annual PAR input of 15 200 mol m -2 , about 95% of incident PAR was absorbed, 50% by leaves, 2 5% by non-foliar tissues and 20% by the ground. The remaining 5% was reflected by the top of the canopy.

Stem biomechanics of three columnar cacti from the Sonoran Desert.

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.

On the mechanical properties of the rare endemic cactus Stenocereus eruca and the related species S. gummosus

We examined the hypothesis that the procumbent growth habit of the rare, columnar cactus Stenocereus eruca is in part the result of a diminution of the mechanical properties of stem tissues by comparing the properties of S. eruca plants with those of the putatively closely related semi-erect shrub S. gummosus. Intact stems and surgically removed anatomically comparable regions of the stems of both species were tested in bending and tension to determine their Youngs modulus and breaking stress. A computer program was used to evaluate the contribution of each region to the capacity of entire stems to resist bending forces. Our analyses indicate that the principal stiffening agent in the stems of both species is a peripheral tissue complex (= epidermis and collenchyma in the primary plant body) that has a significantly higher tensile breaking stress and greater extensibility for S. gummosus than that of S. eruca. Computer simulations indicate that the wood of either species contributes little to bending stiffness, except in very old portions of S. gummosus stems, because of its small volume and central location in the stem. These and other observations are interpreted to support the hypothesis that S. eruca evolved a procumbent growth habit as the result of manifold developmental alterations some of which reduced the capacity of tissues to support the weight of stems.

Does cladode inclination restrict microhabitat distribution for Opuntia puberula (Cactaceae)

In contrast with other Opuntia species, most of the cladodes of Opuntia puberula have a horizontal position. This study explores whether the horizontal cladodes are an adaptive trait to increase light interception in the understory or are a neutral trait, and if this characteristic may prevent its distribution in full sun habitats. Cladode inclination angle and its effect on light interception, cladode temperature, and carbon gain are characterized, and anatomical and physiological traits of upper and lower cladode surfaces are described. Inclination angle was under 50° for 95% of the cladodes, and the frequency of low inclination angles increases as light availability decreases. Nocturnal acid accumulation increased with total daily PPFD intercepted, but no significant differences were detected between typical horizontal cladodes and the few vertical cladodes. Chlorophyll content differed in the upper and lower surfaces of horizontal cladodes; however, chlorenchyma thickness, stomatal conductance, and nocturnal acid accumulation were similar between surfaces. The horizontal position of O. puberula cladodes, which is anatomically determined, restricts it to shaded habitats, where the plants do not overheat, but seems to have no effect on carbon gain.

Functional Diversity in Plants: Implications for Conservation Issues of the Mexican Biodiversity

In this chapter, we explore the functional diversity concept and its importance in several ecological issues, especially maintenance of ecosystem services and conservation. We consider that Mexico’s species megadiversity should be reflected into a high functional diversity. However, our knowledge on this issue is still limited. Interest in the functional diversity approach has just increased in Mexico. Despite that, since the 1970s, ecophysiological research in Mexican ecosystems has had important pioneer contributions to our knowledge on functional traits in plants and its ecological importance. In this chapter, we review some case studies describing our knowledge of plant physiological diversity in different ecosystems, as examples of the high functional diversity in Mexico. Unfortunately anthropogenic disturbance is increasingly affecting species biodiversity, in particular the more vulnerable species and ecosystems. Increasing research on the functional traits of Mexican ecosystems will provide important information about species function at the ecosystem level and species vulnerability in the context of human disturbance and/or climatic change. Studies focused in functional diversity as an important component of biodiversity will provide us a solid base for planning on conservation decisions, restoration programs, and maintenance of ecosystem services.