THE ELASTIC MODULI AND MECHANICS OF POPULUS TREMULOIDES (SALICACEAE) PETIOLES IN BENDING AND TORSION

Changes attending leaf development in the mechanical behavior and elastic moduli of the petioles of Populus tremuloides Michx. were examined in terms of total leaf weight (Wt), lamina and petiole weight (Wl and Wp), petiolar length (L), and lamina surface area (A). A primary concern was the extent to which two elastic moduli (Young's modulus E and the shear modulus G) of petioles changed over time and were correlated with one another. E and G were measured by means of multiple resonance frequency spectra, and together with the dimensions of cross sections through petioles, these two elastic moduli were used to estimate the stiffness of petioles in simple bending and torsion. Petiolar bending was predicted by means of a model incorporating expressions for both the bending stiffness (El) and torsional rigidity (GJ), where I is the second moment of area and J is the torsional constant. The predictions from these models were compared to observed petiolar bendings due to Wl. Additionally, the frequency of the oscillatory motion of leaves (placed in a wind tunnel with a 1 m sec^-1 ambient wind speed, directed normal to the blade of the leaf) was determined. Results indicate that L, A, Wl, Wp, and Wt were positively correlated with the age of leaves (crudely estimated as a function of leaf plastochron index, LPI); these morphometric parameters were also correlated with the magnitudes of E and G. Also, G was positively and linearly correlated with E, and was, on the average, an order of magnitude less than E. EI and GJ were positively correlated with LPI. The relationships among E, G, EI, C and Wl, Wp, Wt are discussed in terms of leaf allometries.     

GRAVITY-INDUCED EFFECTS ON MATERIAL PROPERTIES AND SIZE OF LEAVES ON HORIZONTAL SHOOTS OF ACER SACCHARUM (ACERACEAE)

The influence of gravity on the size and mechanical properties of mature leaves on horizontal shoots and etiolated seedlings of Acer saccharum Marsh. (Aceraceae) was examined. Leaves were grouped into three categories regarding their location on shoots (dorsal or “top” T, lateral or “left/right” L/R, and ventral or “bottom” B). Young's modulus E, petiole length L, lamina surface area A and weight P, and the cross-sectional areas of different tissues within petioles were measured for each leaf and were found to be correlated with leaf location (T, L/R, and B): T leaves were smaller and had lower E than their B counterparts; the size and material properties of L/R leaves were intermediate between those of T and B leaves. In general, A, P, and E decreased from the base to the tip of shoots. In addition to anisophylly, the influence of gravity induced petiole bending and torsion and resulted in the horizontal planation of laminae. This was observed for field-grown mature plants and etiolated seedlings. Petiole bending and torsion were interpreted as gravimorphogenetic phenomena. Anatomically, L, E, and petiole deflection angle Fv measured from the vertical were highly correlated with the combined cross-sectional areas of phloem fibers and xylem in petioles of B leaves and when data from all leaves were pooled. It is tentatively advanced that the correlation of E with the transverse areas of phloem fibers and xylem is evidence that either the pattern or the extent of lignification of petiole tissues is influenced by petiole position with respect to gravity.     

藤本植物Potentilla reptans叶对风和遮阴的响应

生长在一个密集植物群中的植株由于相互遮蔽而不可能对风的影响做出反应,因为这样的环境条件(有限的光资源)对由风导致的矮小表型植株的生长是不利的。为弄清在密集植物群体中生长的植株对风的响应,利用藤本植物Potentilla reptans的10种基因型做实验材料,在温室条件下(光照强度为日光照的50%,红光/远红光=1.2)模拟冠层遮阴(相当于15%的日光照,红光/远红光=0.3),研究了藤本植物叶对风的响应。结果表明,Potentilla reptans的10种基因型植株在冠层遮阴下(低的红光/远红光)都表现出典型的避阴生长响应:较少的叶(叶生物量少),长而细但硬度系数高(higher Young's modulus)的叶柄;而受风影响的植株,无论遮阴或不遮阴,其植株的叶相对较多,叶柄短、粗且柔韧性强(lower Young's modulus),说明Potentilla reptans叶对风的响应并未因遮阴而被压抑,其可塑性变化不过是对复杂生境做出的一种生长权衡:尽可能增强抗风能力(矮壮)和获取最大光能(足够高而避免被遮光),即保证在存活下去的前提下获取最大的生长效率。     

Differences in mechanical and structural properties of surface and aerial petioles of the aquatic plant Nymphaea odorata subsp. tuberosa (Nymphaeaceae)

Lily pads (Nymphaea odorata) exhibit heterophylly where a single plant may have leaves that are submerged, floating, or above (aerial) the surface of the water. Lily pads are placed in a unique situation because each leaf form is exposed to a distinctly different set of mechanical demands. While surface petioles may be loaded in tension under conditions of wind or waves, aerial petioles are loaded in compression because they must support the weight of the lamina. Using standard techniques, we compared the mechanical and morphological properties of both surface and aerial leaf petioles. Structural stiffness (EI) and the second moment of area (I) were higher in aerial petioles, although we detected no differences in other mechanical values (elastic modulus [E], extension ratio, and breaking strength). Morphologically, aerial petioles had a thicker rind, with increased collenchyma tissue and sclereid cell frequency. Aerial petioles also had a larger cross-sectional area and were more elliptical. Thus, subtle changes in the distribution of materials, rather than differences in their makeup, differentiate petiole forms. We suggest that the growth of aerial petioles may be an adaptive response to shading, allowing aerial leaves to rise above a crowded water surface.     

Differences between Acer saccharum Leaves from Open and Wind-Protected Sites

Size-correlated variations of sugar maple (Acer saccharum L.)leaf anatomy and mechanical properties were determined for twosaplings (from open and wind protected sites) to examine theeffects of chronic wind-induced mechanical disturbance on leafsize, rigidity and flexibility. Based on a total sample of 78leaves, comparisons indicated that the mean size of the opensite leaves (n =37) was smaller in every measured respect comparedwith that of the closed site leaves (n =41). Open site leaveshad, on average, smaller lamina surface area, shorter and narrowerpetioles with a smaller volume fraction of lignified tissuesthan those from the closed site. Biomechanical comparisons alsoindicated that the petioles of open site leaves were significantlyless rigid and more flexible than the petioles of closed siteleaves. Despite differences in mean leaf size and petiolar rigidityand flexibility, allometric comparisons indicated the size-dependentvariations in leaf properties were continuous across the twosites. Also, the allocation of leaf biomass with respect tostem biomass along the lengths of the two saplings was statisticallyidentical and indistinguishable from an isometric relationship.However, the smaller diametered branches of the open site saplingbore smaller and fewer leaves with less stiff and rigid petiolesthan those of the closed site sapling. The differences betweenopen and closed site leaves are interpreted to be functionallyadaptive and to indicate that chronic mechanical disturbanceof developingAcer saccharum leaves prefigures mature leaf sizeand petiole properties that have the capacity to reduce winddrag. Results from petioles are contrasted with those of mechanicallydisturbed stems. Leaves; Acer saccharum ; biomechanics; wind-drag; allometry     

Scaling of Petiole Dimensions with Respect to Leaf Size for a Tropical Tree, Scaphium macropodum (Sterculiaceae), in Borneo

Scaphium macropodum (Miq.) Beumee ex Heyne (Sterculiaceae) in a tropical rain forest in West Kalimantan (Indonesia) was analyzed from the viewpoint of statics. The petiole diameter must increase with increasing leaf size to retain enough mechanical stability and a sufficient amount of conductive vessels. The petiole's cross-sectional area at its base was found to be proportional to the leaf blade's dry mass, which indicates that Shinozaki's pipe model is applicable to leaves with different sizes. Although larger leaves produce greater bending moments on the petiole's cross-section as a result of their greater weights, the bending stresses at the petiole's base caused by the leaf's weight were constant at ca. 76,900 g cm⁻² regardless of leaf size. Thicker petioles increase the leaf's mechanical stability, but require sizable energy investments for their construction. It is hypothesized that the constant value for petiolar stress indicates an optimal balance between energy economy and the mechanical stability of S. macropodum leaves. To keep bending stress constant, the leaf blade's center of gravity shifts to a more proximal position and the cross-sectional area of the petiole increases. Received 8 December 1997/ Accepted in revised form 1 December 1998     

Gravity can induce epinastic bending of isolated leaves

Abstract Isolated leaves of Plectranthus fruticosus were grown in cubic plastic cuvettes, and were supplied via their cut petioles with nutrient solution and indole-3-acetic acid (10^?6m ). Holes bored in the cuvette walls allowed the petioles to be oriented at approximately 60°, 90° or 120° to the vertical. Growth of the leaves initially oriented at angles of 60° and 90°, which simulated the situation in the intact plant, did not result in epinastic bending of the petiole. Inversion of the leaves (adaxial surface of the petiole downwards) and orientation of the adaxial/abaxial surfaces of the horizontal petiole parallel to the gravity vector, however, yielded strong epinastic bending of the petioles. In the latter case, this bending was not in the direction of the gravity vector (evidence for point (iii), below). Furthermore, epinastic bending occurred, when the isolated leaves were rotated on a clinostat (petioles parallel to the rotation axis or inclined to the rotation axis at an angle of 30°; 3 r.p.m.). Since a possible influence of the shoot was excluded, it is concluded that (i) perception and response are restricted to the leaf, (ii) gravity alone is sufficient to induce epinasty, (iii) a gravitropic component of the response can probably be excluded. The clinostat induced epinasty may not have been caused by nullifying the effect of gravity but due to continuous gravistimulation of the leaf.     

Genomic fingerprinting of Frankia strains by PCR-based techniques. Assessment of a primer based on the sequence of 16S rRNA gene of Escherichia coli

Submergence stimulates elongation of the leaves of Rumex palustris and under laboratory conditions the maximum final leaf length (of plants up to 7 weeks old) was obtained within a 9 day period. This elongation response, mainly determined by petiole elongation, depends on the availability of storage compounds and developmental stage of a leaf. A starch accumulating tap root and mature leaves and petioles were found to supply elongating leaves with substrates for polysaccharide synthesis in expanding cell walls. Changes in the composition of cell wall polysaccharides of elongated petioles suggest a substantial cell wall metabolism during cell extension. Reduced starch levels or removal of mature leaves caused a substantial limitation of submerged leaf growth. From the 5th leaf onward enough reserves were available to perform submerged leaf growth from early developmental stages. Very young petioles had a limited capacity to elongate. In slightly older petioles submergence resulted in the longest final leaf lengths and these values gradually decreased when submergence was started at more mature developmental stages. Submerged leaf growth is mainly a matter of petiole elongation in which cell elongation has a concurrent synthesis of xylem elements in the vascular tissue. Mature petioles still elongated (when submerged) by cell and tissue elongation only: the annular tracheary elements stretched enabling up to 70% petiole elongation.     

Effects of leaf blade narrowness and petiole length on the light capture efficiency of a shoot

Effects of the length: width ratio of a leaf blade and petiole length on shoot light capture were studied with computer simulation. Both a larger length: width ratio and longer petiole contributed to larger light capture per unit leaf area due to a reduced aggregation of leaf area around the stem. Other conditions being equal, shoots with narrow leaves and no petioles and those with wide leaves with petioles showed similar light capture as long as the mean distance of the leaf blade from the stem was the same. In shoots with a short internode and/or distichous phyllotaxis, however, narrow leaves contributed more to avoiding mutual shading than wide leaves with petioles. The predominance of light coming from a higher angular altitude also favored narrow leaves. The possible consequences of these results in the adaptive geometry of plant architecture are discussed.     

冬性植物红菜薹在不同温度处理下花青素积累的分子机制

芸薹属植物红菜薹（Brassica rapa）是一种常见的蔬菜,它的花茎和叶柄表皮中均积累有花青素。为了解红菜薹中花青素合成的分子机制,进行了花青素含量的测定和花青素合成相关基因的表达分析。研究结果表明,叶柄表皮中的花青素含量显著高于叶片（去主脉）的花青素含量。同时,叶柄表皮花青素合成相关基因的表达水平高于叶柄（去表皮）和叶片（去主脉）的表达水平,这表明红菜薹中花青素的合成调控发生在转录水平。BrMYBA1仅在叶柄表皮中表达,但BrbHLH1和BrWD40在叶片和叶柄表皮中均能检测到表达。因此,BrMYBA1的转录激活可能与红菜薹的花青素合成相关。连续低温处理时,红菜薹叶柄表皮中的花青素含量逐渐增加,而该组织中花青素合成的结构基因表达水平逐渐降低。     

Functional significance of the mandibular symphysis.

The purpose of this investigation was to relate the morphology of connective tissues in the mandibular symphysis to the behavioral and experimental evidence for mobility and mechanical stress at the symphysis. The anatomy of the symphysis was examined histologically in 6 mammalian orders encompassing 22 species. Behavioral and experimental evidence of stress during the power stroke of the chewing cycle correspond with stresses at the symphysis implied by the location and orientation of symphyseal connective tissues. These stresses are: (1) dorsoventral shear of the symphysis due to the transfer of force from balancing to chewing sides, (2) bending of the symphysis causing tension along the inferior and compression along superior borders due to torsion on the dentaries from the jaw closing muscles, and (3) antero-posterior shear of the symphysis due to an anteriorly directed stress on the chewing side. Interspecific comparisons suggest that leaf eaters can resist greater dorsoventral shear than fruit or insect eaters, but no correlations exist between diet and bending or antero-posterior shear. This suggests that chewing leaves requires larger biting forces.     

Petiole mechanics, leaf inclination, morphology, and investment in support in relation to light availability in the canopy of Liriodendron tulipifera

To determine the role of leaf mechanical properties in altering foliar inclination angles, and the nutrient and carbon costs of specific foliar angle variation patterns along the canopy, leaf structural and biomechanical characteristics, biomass partitioning into support, and foliar nitrogen and carbon concentrations were studied in the temperate deciduous species Liriodendron tulipifera L., which possesses large leaves on long petioles. We used beam theory to model leaf lamina as a uniform load, and estimated both the lamina and petiole flexural stiffness, which characterizes the resistance to bending of foliar elements at a common load and length. Petiole and lamina vertical inclination angles with respect to horizontal increased with increasing average daily integrated photon flux density (Q_int). Yet, the light effects on lamina inclination angle were primary determined by the petiole inclination angle. Although the petioles and laminas became longer, and the lamina loads increased with increasing Q_int, the flexural stiffness of both lamina and petiole increased to compensate for this, such that the lamina vertical displacement was only weakly related to Q_int. In addition, increases and decreases in the petiole inclination angle with respect to the horizontal effectively reduced the distance of lamina load from the axis of rotation, thereby reducing the bending moments and lamina inclination due to gravity. We demonstrate that large investments, up to 30% of total leaf biomass, in petiole and large veins are necessary to maintain the lamina at a specific position, but also that light has no direct effect on the fractional biomass investment in support. However, we provide evidence that apart from light availability, structural and chemical characteristics of the foliage may also be affected by water stress, magnitude of which scales positively with Q_int.     

Effects of pod removal on the transport and accumulation of abscisic Acid and indole-3-acetic Acid in soybean leaves

Concentrations of abscisic acid (ABA) and indole-3-acetic acid (IAA) in the second most recently expanded trifoliolate leaf were determined during reproductive development of soybean (Glycine max [L.] Merr cv `Chippewa 64'). The concentration of ABA in leaves was constant during most of the seed filling period until the seeds began to dry. The concentration of IAA in the leaves decreased throughout development. Removal of pods 36 hours prior to sampling resulted in increased concentrations of ABA in leaves during the period of rapid pod filling but had little effect on the concentration of IAA in leaves. ABA appears to accumulate in leaves after fruit removal only when fruits represent the major sink for photosynthate.

ABA and IAA moving acropetally and basipetally in petioles of soybean were estimated using a phloem exudation technique. ABA was found to move mostly in the basipetal direction in petioles (away from laminae). IAA, primarily in the form of ester conjugate(s), was found to be moving acropetally (toward laminae) in petioles. The highest amount of IAA ester(s) was found in petiole exudate during the mid and late stages of seed filling. Removal of fruits 36 hours prior to exudation reduced the amount of IAA ester recovered in exudate, suggesting that fruits were a source of the IAA conjugate in petiole exudate.

     

Are leaves ‘freewheelin'? Testing for a Wheeler‐type effect in leaf xylem hydraulic decline

A recent study found that cutting shoots under water while xylem was under tension (which has been the standard protocol for the past few decades) could produce artefactual embolisms inside the xylem, overestimating hydraulic vulnerability relative to shoots cut under water after relaxing xylem tension (Wheeler et al. 2013). That study also raised the possibility that such a ‘Wheeler effect’ might occur in studies of leaf hydraulic vulnerability. We tested for such an effect for four species by applying a modified vacuum pump method to leaves with minor veins severed, to construct leaf xylem hydraulic vulnerability curves. We tested for an impact on leaf xylem hydraulic conductance (K_x) of cutting the petiole and minor veins under water for dehydrated leaves with xylem under tension compared with dehydrated leaves after previously relaxing xylem tension. Our results showed no significant ‘cutting artefact’ for leaf xylem. The lack of an effect for leaves could not be explained by narrower or shorter xylem conduits, and may be due to lesser mechanical stress imposed when cutting leaf petioles, and/or to rapid refilling of emboli in petioles. These findings provide the first validation of previous measurements of leaf hydraulic vulnerability against this potential artefact.     

Petiole twisting in the crowns of<Emphasis Type="Italic"> Psychotria limonensis:</Emphasis> implications for light interception and daily carbon gain

We used Y-plant, a computer-based model of crown architecture, to examine the implications of leaf reorientation resulting from petiole bending in Psychotria limonensis (Rubiaceae) seedlings. During this reorientation process, bending of the petioles of lower leaves that are potentially self-shaded by the upper leaves rotates the lamina around the stem's orthotropic axis so that self-shading is reduced. Simulations of daily light capture and assimilation revealed a 66% increase in daily C gain due to reorientation of the leaves as compared to simulations where the leaves remained in their characteristic opposite decussate pattern set by the phyllotaxy. This was due to enhanced carbon (C) gain of the lower leaves because of the reduction of shading by upper developing leaves in the same vertical plane. The light signal for this movement was experimentally examined by placing leaf-shaped filters above already fully expanded leaves and following the resulting shade-avoiding movements. The filters were either neutral density shade cloth that reduced the photon flux density (PFD) but did not alter the red to far red ratio (R:FR) or a film that reduced the PFD equivalently but also reduced the R:FR. Leaf reorientation was much more rapid and complete under the low R:FR as compared to the high R:FR indicating involvement of a phytochrome photosensory system that detected the presence of a shading leaf. Plants in gaps were found to lack a reorientation response indicating that the reorientation is specific to the shaded understory environment.     

A mechanical perspective on foliage leaf form and function

The mechanical behaviour of large foliage leaves in response to static and dynamic mechanical forces is reviewed in the context of a few basic engineering principles and illustrated in terms of species drawn from a variety of vascular plant lineages. When loaded under their own weight or subjected to externally applied forces, petioles simultaneously bend and twist, and thus mechanically operate as cantilevered beams. The stresses that develop in petioles reach their maximum intensities either at their surface or very near their centroid axes, where they are accommodated either by living and hydrostatic tissues (parenchyma and collenchyma) or dead and stiff tissues (sclerenchyma and vascular fibres) depending on the size of the leaf and the species from which it is drawn. Allometric analyses of diverse species indicate size-dependent variations in petiole length, transverse shape, geometry and stiffness that accord well with those required to maintain a uniform tip-deflection for leaves with laminae differing in mass. When dynamically loaded, the laminae of many broad-leaved species fold and curl into streamlined objects, thereby reducing the drag forces that they experience and transmit to their subtending petioles and stems. From a mechanical perspective, the laminae of these species operate as stress-skin panels that distribute point loads more or less equally over their entire surface. Although comparatively little is known about the mechanical structure and behaviour of foliage leaves, new advances in engineering theory and computer analyses reveal these organs to be far more complex than previously thought. For example, finite-element analyses of the base of palm leaves reveal that stresses are decreased when these structures are composed of anisotropic as opposed to isotropic materials (tissues).     

Paths and Mechanisms of Moisture Movements in Detached Leaves of White Clover(Trifolium repens L.): I. Losses of Petiole Moisture direct from Petioles and via Laminae

The experiments described were concerned with losses of moisturefrom laminae and petioles of leaves of white clover (Trifoliumrepens L.) detached from parent plants, and with the resistancesin the two pathways of loss of petiole moisture-via laminaeand direct from petiole surfaces. Methods involving determination of moisture losses from separatedlaminae and petioles were unsatisfactory and transpiration chamberswere therefore used to permit determinations with intact leaves. Up to 24 per cent of the total petiole moisture was lost viathe laminae, the amount frequently exceeding that initiallyresiding in the xylem of the petioles. Stomatal, cuticular, and external resistances were calculatedat several stages of wilting. Rates of loss of petiole moistureby both pathways were controlled by stomatal and external resistancesin the early stages and by cuticular and internal resistancesin the later stages. Under certain conditions, transpirationof petiole moisture via the lamina ceased earlier than mighthave been expected; estimates were made of the resistances involved.     

Effects of Wind on the Allometry of Two Species of Plants in an Elfin Cloud Forest

Thigmomorphogenesis includes the effects of mechanical perturbation on plant growth. To test whether thigmomorphogenesis is evident at different scales within plants, we investigated the effect of wind on allometric relationships between specific plant parts. We chose two species from the elfin cloud forest of Puerto Rico that have contrasting growth habits, the shrub Clibadiun erosum (Asteraceae) and the palm Prestoea acuminata var. montana (Arecaceae), and subjected them to barrier-protected and wind-exposed treatments. For C. erosum , we compared the allometry of stems and branches against three allometric models that predict that plant height or branch length increases at the 1, 2/3, and 1/2 power of stem diameter. Only the geometric similarity model (scaling exponent of 1) seemed to hold when plants were exposed to the wind. We found relatively fewer leaves per number of branches produced and fewer leaves per increment of branch diameter in the plants of C. erosum exposed to the wind. Mean petiole length ratios (petiole length/basal radius) of P. acuminata were higher on leaves of barrier-protected plants for both simple and compound leaves, indicating that petioles were stouter and mechanically safer in the wind-exposed plants. We suggest that alteration of the allometric relationships of plant parts, organs, or plant modules (stems and branches of C. erosum and leaves of P. acuminata ) and alteration of the number of plant parts (leaves and branches of C. erosum ) are adaptive responses resulting from the mechanical perturbation induced by wind in the elfin forest.     

The twist-to-bend compliance of the Rheum rhabarbarum petiole: integrated computations and experiments

Plant petioles can be considered as hierarchical cellular structures, displaying geometric features defined at multiple length scales. Their macroscopic mechanical properties are the cumulative outcome of structural properties attained at each level of the structural hierarchy. This work appraises the compliance of a rhubarb stalk by determining the stalk’s bending and torsional stiffness both computationally and experimentally. In our model, the irregular cross-sectional shape of the petiole and the layers of the constituent tissues are considered to evaluate the stiffness properties at the structural level. The arbitrary shape contour of the petiole is generated with reasonable accuracy by the Gielis superformula. The stiffness and architecture of the constituent layered tissues are modeled by using the concept of shape transformers so as to obtain the computational twist-to-bend ratio for the petiole. The rhubarb stalk exhibits a ratio of flexural to torsional stiffness 4.04 (computational) and 3.83 (experimental) in comparison with 1.5 for isotropic, incompressible, circular cylinders, values that demonstrate the relative structural compliance to flexure and torsion.