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     

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).     

Leaf recruitment and elongation: an adaptive response to flooding in Villarsia reniformis

Villarsia reniformis (Menyanthaceae) responds to flooding by rapid leaf elongation and continual recruitment of young, submerged leaves (4.3–6.5 per week). Leaf production is influenced by nutrient availability and water depth. Leaves are submerged and die as the water level rises, but are replaced by younger leaves able to broach the surface. Young petioles may elongate at more than 10 cm per day, but lose the ability to elongate after the blades are exposed to air more than twice. Young petioles produce new cells and existing cells elongate, but in older petioles fewer new cells are produced and cell elongation, whilst limited, is the main mechanism for petiole elongation. Continual recruitment implies a high cost for production of structural tissue, but ensures that leaves capable of rapid extension are within reach of the water surface and the plants can respond quickly to flooding.     

Scaling sun and shade photosynthetic acclimation of Alocasia macrorrhiza to whole-plant performance – I. Carbon balance and allocation at different daily photon flux densities

We determined the carbon allocation patterns and construction costs of Alocasia macrorrhiza plants grown at different photon flux densities (PFD) as well as the whole-plant carbon gain of these plants at different daily PFDs. Growth at high PFD resulted in thicker leaves with a higher leaf mass per unit area, and increased biomass allocation to petioles and roots, as compared to growth at low PFD. Increased allocation to petioles may have been necessary to support the heavier leaves, whereas increased allocation to roots may have been necessary to supply sufficient water for the higher transpiration rates in high PFD. Root biomass was highly correlated with the daily, whole-plant transpiration rate. Tissue construction costs per unit dry mass were unchanged by acclimation, but, since the mass per unit areas of leaves, roots and petioles all increased, construction costs per unit leaf area were much higher for plants grown at high PFD. On a per unit leaf area basis, daily whole-plant carbon gain measured at high daily PFD was higher in high- than in low-PFD-grown plants. However, on a per unit leaf mass basis, low-PFD-grown plants had a daily carbon gain at least as high as that of high-PFD-grown plants at high daily PFD. At low daily PFD, low-PFD-grown plants maintained an advantage over high-PFD-grown plants in terms of carbon gain because of their larger leaf area ratios. Thus, in terms of carbon gain, low-PFD-grown plants performed better than sun plants at low PFD and as well as high-PFD-grown plants at high PFD, despite their lower photosynthetic capacities per unit area. For high-PFD-grown plants, the higher construction costs per unit leaf area resulted in lower leaf area ratios, which counteracted the advantage of higher photosynthetic rates per unit leaf area.     

Availability of water and inorganic nutrients in the persistent leaf bases of the grasstree Kingia australis, and uptake and translocation of labelled phosphate by the embedded aerial roots

Numerous lateral branches of the concealed aerial roots of the grasstree Kingia australis ramify through the persistent leaf bases, suggesting a role in uptake of water and nutrients localed up to 8 m above ground. These leaf bases were shown to hold up to more than three times their weight in water. Water and nutrients (N, P and K) available in the leaf bases may greatly exceed that in an equal volume of surface soil, especially during the dry summer months. At this time, ³²P injected# among the leaf bases is strongly absorbed by the aerial roots and translocated to the meristematic regions of other laterals beneath the point of application, but mostly to the stern and leaves above it. Scattered vascular bundles in the pith are the major routes for the rapid distribution of "P up and down the trunk following its departure from the aerial roots. Water and nutrients stored in the leaf bases and their subsequent uptake by the associated aerial roots may therefore contribute significantly to this species' tolerance of long summer droughts and extremely impoverished soils.     

Anatomical and morphological parameters of leaves and leaf petioles of Actinidia deliciosa

Differences in anatomy and morphology of the kiwifruit leaves and leaf petioles might play a considerable role in the sex-determination. Three months after bud break (June), the kiwifruit leaves of both male and female plants, grown on the vegetative and generative shoots showed different leaf area (128.6 ± 13.45 cm2 in male and 104.5 ± 4.02 cm2 in female plants) and shape. The most frequently leaf shape was determined as "folium cordatum" and "folium rotundato-cordatum". Higher values of total leaf thickness of the female leaves (190 ± 3.84 μm) in comparison to male leaves (174 ± 3.52 μm) were estimated, resulting in the thicker adaxial leaf epidermis and especially in thicker palisade parenchyma in female leaves (136 ± 2.76 μm in comparison to 104 ± 1.61 μm in male leaves). Typically bifacial leaves were observed in both male and female leaves. Anomocytic stomata in hypostomatic leaves were found. The reticulate venation appears to be the main type of leaf venation. Stalked stellate multicellular trichomes on the abaxial leaf side were frequently observed in the leaves of both sexes. No important differences between male and female plants were found in the structures of vascular system in leaves and leaf petioles. Thus leaf thickness and surface morphology of adaxial leaf epidermis can be considered as important structural parameters in the sex determination. This revised version was published online in July 2006 with corrections to the Cover Date.     

含羞草(Mimosa pudica L.)叶片匀浆悬浮液的滞后环

由于受力后叶子立即发生运动 ,含羞草是一个研究力对于生物细胞作用的良好模型。在以往的研究中 ,人们认为此种现象与受力后渗透压改变、离子通道被激活、细胞骨架的动态变化有关。该文旨在通过观察含羞草叶片和叶柄匀浆悬浮液的应力 切变率滞后环变化 ,揭示含羞草的力学性质。在用于比较的含羞草、叶下珠和猪骨骼肌匀浆悬浮液以及水 4个系统中 ,只有含羞草系统具有明显的逆时针滞后环轨迹 ,而其它的 3个系统均不存在。以上结果提示 ,在含羞草的匀浆悬浮液系统中 ,有一种或多种物质 (可能是蛋白质和细胞骨架 )在剪切应力作用过程中由颗粒状结构向网状结构转变 ,由无序结构向有序结构转变 ,由液体结构向黏弹性状态转变 ,而当力撤除以后再缓慢恢复。     

EFFECTS OF TISSUE VOLUME AND LOCATION ON THE MECHANICAL CONSEQUENCES OF DEHYDRATION OF PETIOLES

Data are presented on the mechanical consequences of dehydration for the petioles of two monocots and two dicots differing in leaf morphology (pinnate leaves ofChamaedorea erumpens and simple leaves of Spathiphyllum ‘clevelandii‘; pinnate leaves of Acer negundo and simple leaves of A. saccharum). The flexural stiffness EI of petioles decreased over a broad range of tissue water potential (– 10 < ψ_w <– 50 bars). Within the same range of ψ, the second moment of area I and the elastic modulus E were observed to decrease and increase, respectively. However, the mechanical alterations of Chamaedorea and A. negundo petioles were significantly less than those observed for Spathiphyllum and A. saccharum petioles. The increase in E of Spathiphyllum and A. saccharum petioles attending dehydration was linearly correlated with an increase in the relative volume fraction of tissues with lignified, thick cell walls (“support tissues”). The decrease in I of Spathiphyllum and A. saccharum petioles was linearly correlated with a decrease in the relative volume fraction of tissues with nonlignified, thin cell walls (“ground tissues”). Similar trends were observed for the petioles of C. erumpens and A. negundo but were found not to be statistically significant. Anatomical differences in the relative volume fraction and spatial locations of support tissues in the petioles of these four taxa appear to account for the differences observed in the mechanical consequences of petiole dehydration.     

Reduction of auxin transport capacity with age and internal water deficits in cotton petioles

Auxin transport was examined in leaf petioles taken from the upper, middle, and lower leaf canopy of large cotton plants. The ability of petioles to transport auxin decreased with age (position) of the leaves. Plant water deficit reduced transport regardless of age. These correlations support the view that reduced transport capacity of petioles plays a significant role in the induction of abscission of lower or older leaves during water deficits.     

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.     

水淹导致皇冠草光合机构发生变化并加剧其出水后光抑制

通过气体交换和叶绿素荧光等方法研究了水淹及胁迫解除后皇冠草不同功能叶的光合特性及光抑制的变化.结果表明:与对照相比,气生叶(全淹组淹水前形成的功能叶)在水淹条件下叶片大小和气孔没有明显变化,但沉水叶(全淹组淹水后新生的功能叶)的叶面积增加,气孔变小,上表皮气孔密度增加.水淹导致气生叶碳同化能力、光化学效率和叶绿素含量下降.沉水叶在发育过程中碳同化能力、光化学效率和叶绿素逐渐升高.气生叶和沉水叶出水后其活体叶片在强光下的相对含水量急剧下降,发生明显的光抑制;而弱光下无明显光抑制发生.出水后离体叶片强光照射下6h后两种功能叶均发生严重光抑制,且弱光下不能恢复.因此,可以认为淹水条件下,沉水叶上表皮气孔密度的增加使其蒸腾速率提高;沉水叶较强的碳同化能力和增加的叶面积是确保其植株水下生存的重要因素;强光使气生叶和沉水叶出水后均发生严重光抑制,导度和蒸腾速率提高导致的叶片失水则加剧了这一过程,两者共同作用导致自然条件下两种功能叶的出水死亡.     

Climatic classification and ordination of the Spanish Sistema Central: relationships with potential vegetation

Comparisons among European, Japanese and North-American temperate deciduous woody floras revealed that there is no difference in shade-tolerance or in successional position between the compound- and simple-leaved species. Given that the compound-leaved species usually have greater biomass investments in non-productive throwaway supporting structures, it remained unclear how they could be as shade-tolerant as the simple-leaved analogues. To find out the role of the variability in leaf structure and composition in shade-tolerance of these species, foliar morphology and chemistry were analysed in 15 Estonian temperate compound-leaved deciduous woody taxa.Both foliar morphological and chemical parameters influenced the fractional investment of foliar biomass in petioles. The proportion of leaf biomass in petioles was independent of leaf size, but it increased with increasing leaflet number per leaf, suggesting that spacing rather than support requirements determined the biomass investment in petioles. The leaves with greater nitrogen concentrations also had larger foliar biomass investments in petioles. The latter effect possibly resulted from a greater water demand of functionally more active protein-rich leaves. Though the proportion of leaf biomass invested in petioles was high (for the whole material on average 15.9±0.4%), petioles were considerably cheaper to construct in terms of mineral nutrients than leaflets. e.g., petioles contained on average only 5.55±0.14% of total leaf nitrogen. Since in many cases the availability of mineral nutrients such as nitrogen rather than organic carbon sets limits to total leaf biomass on the plant, I suggested, contrary to previous claims, that the costs for foliage formation should not necessarily be different between compound- and simple-leaved species. Compound-leaved species also fit the basic relationships previously observed in simple-leaved analogues. Leaf size increased and leaf dry mass per area (LMA) decreased with increasing shade-tolerance. Thus, more shade-tolerant species construct a more effective foliar display for light interception at low irradiance with similar biomass investment in leaves. Species shade-tolerance was independent of biomass investment in petioles. However, due to the genotypic plasticity in LMA, more shade-tolerant species supported more foliar area at a constant leaf biomass investment in petioles.     

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.     

Occurrence of endodermis with a casparian strip in stem and leaf

It is well known that an endodermis with casparian strip always occurs in roots, but few people are aware that it also occurs in stems and leaves of some vascular plants. The rather sparse literature on endodermis in aerial organs was last included in a review in 1943. The present compilation, which does not consider hydathodes, nectaries, or other secretory structures, emphasizes distribution of cauline and foliar endodermis with casparian strip. It occurs unevenly among major taxa: quite common in rhizomes and leaves among pteridophyte groups, with exceptions; absent in gymnosperm stems but found in leaves at least among some conifers; in stems of at least 30 mostly herbaceous angiosperm families, but far less common in leaves, where it is mostly reported from petioles. Etiolation can induce casparian strips in stems and petioles of some herbaceous plants, but results from leaf blades are questionable. There are recent reports of an endodermis with casparian strip in leaves of both woody and herbaceous taxa. The physiological function, if any, of a casparian strip in aerial organs remains unknown.     

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.     

Effects of light quality on leaf morphogenesis of a heterophyllous amphibious plant, Rotala hippuris

Background and Aims

For heterophyllous amphibious plants that experience fluctuating water levels, it is critical to control leaf development precisely in response to environmental cues that can serve as a quantitative index of water depth. Light quality can serve as such a cue because the ratio of red light relative to far-red light (R/FR) increases and blue-light intensity decreases with increasing water depth. Growth experiments were conducted to examine how R/FR and blue-light intensity alter leaf morphology of a heterophyllous amphibious plant, Rotala hippuris.

Methods

Using combinations of far red (730 nm), red (660 nm) and blue (470 nm) light-emitting diodes (LEDs), growth experiments were used to quantitatively evaluate the effects of the R/FR ratio and blue-light intensity on leaf morphology.

Key Results

Under the natural light regime in an outside growth garden, R. hippuris produced distinct leaves under submerged and aerial conditions. R/FR and blue-light intensity were found to markedly affect heterophyllous leaf formation. Higher and lower R/FR caused leaf characters more typical of submerged and aerial leaves, respectively, in both aerial and submerged conditions, in accordance with natural distribution of leaf types and light under water. High blue light caused a shift of trait values toward those of typical aerial leaves, and the response was most prominent under conditions of R/FR that were expected near the water surface.

Conclusions

R/FR and blue-light intensity provides quantitative cues for R. hippuris to detect water depth and determine the developmental fates of leaves, especially near the water surface. The utilization of these quantitative cues is expected to be important in habitats where plants experience water-level fluctuation.     

The Comparative Biology of Closely Related Species: VI. ANALYSIS OF THE GROWTH OF TRIFOLIUM REPENS AND T. FRAGIFERUM IN PURE AND MIXED POPULATIONS

An analysis was made of the first season's growth of pure andmixed stands of Trifolium repens and T. fragiferum. T. repens,although having smaller seeds and cotyledon area, achieved afaster early rate of leaf production than T. fragiferum andquickly developed a larger area of leaf both in pure standsand in mixtures. By the 18th and 21st weeks from sowing, T.fragiferum had more elongated petioles than T. repens, leaveswere borne higher in the canopy and contributed an increasingpart to the Leaf Area Index of mixtures. The increase in leaf area of swards with time was largely associatedwith increased area of individual leaves. The differences betweenspecies were largely due to differences in number of leaves. These results are discussed in relation to the conditions underwhich stable associations of species may be formed.     

Changes in Stem and Leaf Hydraulics Preceding Leaf Shedding in Castanea Sativa L.

This paper describes changes in leaf water status and in stem, petiole and leaf blade hydraulics preceding leaf senescence and shedding in Castanea sativa L. (chestnut). Measurements of maximum diurnal leaf conductance to water vapour (g_L), minimum water potential (_L), hydraulic conductance per unit leaf surface area of stems (K_SL), petioles (K_PL) and leaf blades (K_LL) and number of functional conduits and inside diameter distribution were performed in June, September and October 1999. In September, still green leaves had undergone some dehydration as indicated by decreased g_L (by 75 %) and _L with respect to June. In the same time, K_SL decreased by 88 %, while K_PL and K_LL decreased by 50 % and 20 % of the conduits of stems and 10 % of the petioles (all belonging to the widest diameter range) were no longer functioning, causing a decrease in the theoretical flow by 82 % in stems and 27 % in petioles. Stem xylem blockage was apparently due to tyloses growing into conduits. We advance the hypothesis that the entire process of leaf shedding and winter rest may be initiated by extensive stem embolism occurring during the summer.     

LIGHT EFFECTS ON LEAF MORPHOLOGY IN WATER HYACINTH (EICHHORNIA CRASSIPES)

Water hyacinth leaves in natural populations vary from being long and thin-petioled to being short with inflated petioles. A variety of factors has been used experimentally to alter water hyacinth leaf shape, but what controls the development of leaf morphology in the field has not been established. We measured photosynthetic photon flux density (PPFD) and spectral distribution of radiation in a natural water hyacinth population. PPFD in the center of the water hyacinth mat was reduced to 2.7% of full sunlight, and the red to far red (R:FR) ratio was reduced to 0.28. When shoot tips of plants were exposed to artificial light environments, only plants in the treatment with a R:FR ratio comparable to that in the natural population produced leaves with long, thin petioles. Shoot tips in full sun or covered with clear plastic bags or bags that reduced light quantity without greatly altering light quality produced shorter leaves with inflated petioles. We hypothesize that the altered light quality inside a mat is a major environmental control of water hyacinth leaf morphology.     

The structure of xylem vessels in grapevine (Vitaceae) and a possible passive mechanism for the systemic spread of bacterial disease

Xylem-dwelling pathogens become systemic, suggesting that microorganisms move efficiently in the xylem. To better understand xylem pathways and how bacteria move within the xylem, vessel connectivity between stems and leaves of Vitis vinifera cv. Chardonnay and Muscadinia rotundifolia cv. Cowart was studied. Three methods were used: (1) the light-producing bacterium, Yersinia enterocolitica, (Ye) strain GY5232 was loaded into petioles and followed using X-ray film, (2) fluorescent beads were loaded and followed by microscopy, and (3) low-pressure air was pumped into leaves and extruded bubbles from cuts in submerged leaves were followed. Bacteria, beads, and air moved through long and branched xylem vessels from the petiole into the veins in leaves of both varieties. From the stem, bacteria and air traveled into primary and secondary veins of leaves one, two, and three nodes above the loading point of the bacteria or air. Particles and air could move unimpeded through single xylem vessels or multiple vessels (conduits) connected possibly through broken pit membranes from within the stem axis into leaf blades. Bacteria were also able to move long distances within minutes from stem to leaf passively without having to cross pit membranes. Such complex, open xylem conduits have not been well documented before; these findings will help elucidate mechanisms involved in the systemic spread of pathogens.