How to become a tree without wood – biomechanical analysis of the stem of Carica papaya L

Carica papaya L. does not contain wood, according to the botanical definition of wood as lignified secondary xylem. Despite its parenchymatous secondary xylem, these plants are able to grow up to 10‐m high. This is surprising, as wooden structural elements are the ubiquitous strategy for supporting height growth in plants. Proposed possible alternative principles to explain the compensation for lack of wood in C. papaya are turgor pressure of the parenchyma, lignified phloem fibres in the bark, or a combination of the two. Interestingly, lignified tissue comprises only 5–8% of the entire stem mass. Furthermore, the phloem fibres do not form a compact tube enclosing the xylem, but instead form a mesh tubular structure. To investigate the mechanism of papaya's unusually high mechanical strength, a set of mechanical measurements were undertaken on whole stems and tissue sections of secondary phloem and xylem. The structural Young's modulus of mature stems reached 2.5 GPa. Since this is low compared to woody plants, the flexural rigidity of papaya stem construction may mainly be based on a higher second moment of inertia. Additionally, stem turgor pressure was determined indirectly by immersing specimens in sucrose solutions of different osmolalities, followed by mechanical tests; turgor pressure was between 0.82 and 1.25 MPa, indicating that turgor is essential for flexural rigidity of the entire stem.     

Hygrochastic capsule dehiscence in New Zealand alpine Veronica (Plantaginaceae)

? Premise of the study: Plant movement is more widespread than often recognized, involving different organs and mechanisms. Hygrochasy (opening in response to moisture) is a capsule-opening movement that is widely believed to be predominantly a feature of plants of desert and arid zones, where it may protect against seed predators and harsh climatic conditions and restrict dispersal to favorable germination times and sites. However, recently it has been reported from a wider range of environments. This study demonstrates that hygrochasy is much more common than previously realized, extending the phenomenon to plants of alpine habitats. ? Methods: Capsules of 23 species of New Zealand Veronica were collected, and we used light microscopy to investigate the anatomy and biomechanics responsible for the opening mechanism. Additionally, we collected morphological data to identify common traits of hygrochastic species. ? Key results: Hygrochastic capsule dehiscence was found in 10 alpine Veronica species. The opening mechanism is based on an antagonistic reaction between a nonlignified swelling tissue and a lignified resistance tissue. In Veronica, hygrochasy is associated with erect, narrowly angustiseptate capsules on short peduncles of creeping subshrubs or cushion plants. ? Conclusion: Hygrochasy is a common dehiscence type in New Zealand alpine Veronica, and for the first time, this mechanism is described in detail for plants in alpine habitats. We propose that hygrochasy provides an effective seed dispersal mechanism in solitary capsules embedded in cushion plants and may restrict dispersal within habitat patches.     

Compressive/tensile stresses and lignified cells as resistance components in joints between cladodes of Opuntia laevis (Cactaceae): responses to applied stresses

Cactaceae are a diverse group of plants with a wide variety of morphologies and reproductive strategies. Many species have segmented stems in which terminal cladodes may be separated from main stem cladodes with varying amounts of resistance. Previous results demonstrated that lignified xylem cells in tensile portions of stem joints provide the main resistance to separation of cladodes within cactus plants. The purpose of the present study was to determine if stem joints of Opuntia laevis would produce additional lignified xylem cells in response to additional externally applied stresses. Normal average stress levels, which accompany the addition of a new cladode, were applied to 12 plants. In contrast, double the average stress levels were applied to 13 other plants. After exposure to the two stress regimens for 6 months, the amount and location of lignified xylem cells in joint segments were similar for both stress treatments. So, although the results support the hypothesis that lignified xylem cells act as the main resistance to stress at joints of cladodes, doubling the normal amount of applied stress was insufficient to alter the amount or location of lignified xylem cells in stem joints. These results indicate that normal amounts of lignified xylem cells can resist up to two times the normal amount of stress for 6 months without producing additional lignified xylem cells.     

Gravitropism of cucumber hypocotyls: biophysical mechanism of altered growth

Abstract. The biophysical basis for the changes in cell elongation rate during gravitropism was examined in aetiolated cucumber ( Cucumis sativus L.) hypocotyls. Bulk osmotic pressures on the two sides of the stem and in the epidermal cells were not altered during the early time course of gravitropism. By the pressureprobe technique, a small increase in turgor (0.3 bar, 30 kPa) was detected on the upper (inhibited) side, whereas there was a negligible decrease in turgor on the lower (stimulated) side. These small changes in turgor and water potential appeared to be indirect, passive consequences of the altered growth and the small resistance for water movement from the xylem, and indicated that the change in growth was principally due to changes in wall properties. The results indicate that the hydraulic conductance of the watertransport pathway was large (.25 h ¹ bar ¹) and the water potential difference supporting cell expansion was no greater than 0.3 bar (30 kPa). From pressureblock experiments, it appeared that upon gravitropic stimulation (1) the yield threshold of the lower half of the stem did not decrease and (2) the wall on the upper side of the stem was not made more rigid by a cross-linking process. Mechanical measurements of the stress/strain properties of the walls showed that the initial development of gravitropism did not involve an alteration of the mechanical behaviour of the isolated walls. Thus, gravitropism in cucumber hypocotyls occurs principally by an alteration of the wall relaxation process, without a necessary change in wall mechanical properties.     

The dynamic pipeline: hydraulic capacitance and xylem hydraulic safety in four tall conifer species

Recent work has suggested that plants differ in their relative reliance on structural avoidance of embolism versus maintenance of the xylem water column through dynamic traits such as capacitance, but we still know little about how and why species differ along this continuum. It is even less clear how or if different parts of a plant vary along this spectrum. Here we examined how traits such as hydraulic conductivity or conductance, xylem vulnerability curves, and capacitance differ in trunks, large‐ and small‐diameter branches, and foliated shoots of four species of co‐occurring conifers. We found striking similarities among species in most traits, but large differences among plant parts. Vulnerability to embolism was high in shoots, low in small‐ and large‐diameter branches, and high again in the trunks. Safety margins, defined as the pressure causing 50% loss of hydraulic conductivity or conductance minus the midday water potential, were large in small‐diameter branches, small in trunks and negative in shoots. Sapwood capacitance increased with stem diameter, and was correlated with stem vulnerability, wood density and latewood proportion. Capacitive release of water is a dynamic aspect of plant hydraulics that is integral to maintenance of long‐distance water transport.     

Distribution of vessel size,vessel density and xylem conducting efficiency within a crown of silver birch (<Emphasis Type="Italic">Betula pendula</Emphasis>)

Spatial patterns in vessel diameter, vessel density and xylem conducting efficiency within a crown were examined in closed-canopy trees of silver birch (Betula pendula). The variation in anatomical and hydraulic characteristics of branches was considered from three perspectives: vertically within a crown (lower, middle and upper crown), radially along main branches (proximal, middle and distal part), and with respect to branch orders (first-, second- and third-order branches). Hydraulically weighted mean diameter of vessels (D _h) and theoretical specific conductivity of the xylem (k _t) exhibited no vertical trend within the tree crown, whereas leaf-specific conductivity of the xylem (LSC_t) decreased acropetally. Variation in LSC_t was governed by sapwood area to leaf area ratio (Huber value) rather than by changes in xylem anatomy. The acropetal increase in soil-to-leaf conductance (G _T) within the birch canopy is attributable to longer path length within the lower-crown branches and higher hydraulic resistance of the shade leaves. D _h, k _t and LSC_t decreased, while vessel density (VD) and relative area of vessel lumina (VA) increased distally along main branches. A strong negative relationship between vessel diameter and VD implies a trade-off between hydraulic efficiency and mechanical stability of xylem. D _h and VD combined explained 85.4% of the total variation of k _t in the regression model applied to the whole data set. Xylem in fast-growing branches (primary branches) had greater area of vessel lumina per unit cross-sectional area of sapwood, resulting in a positive relationship between branch radial growth rate and k _t. D _h, k _t and LSC_t decreased, whereas VD increased with increasing branch order. This pattern promotes the hydraulic dominance of primary branches over the secondary branches and their dominance over tertiary branches. In this way crown architecture contributes to preferential water flow along the main axes, potentially providing better water supply for the branch apical bud and foliage located in the outer, better-insolated part of the crown.     

STEM ANATOMY AND ASPECTS OF DEVELOPMENT IN TOMATO

Aspects of anatomical development were correlated with internodal growth in tomato plants, variety ‘Yellow Plum,’ grown for more than 3 months. Internodal length was measured weekly in control plants and those harvested for anatomical study. Gross structure indicated progressive development with increasing age. Primary xylem and phloem first mature in distinct strands and the strands are joined laterally by procambium to form a continuous vascular cylinder. Primary phloem occurs on the outer periphery of the procambium between the early-formed vascular strands. Successive periclinal divisions in the procambium during internode elongation give rise to pronounced radial seriations of the cells. Procambial derivatives are included in the cylinder of thick-walled, lignified vascular cells that become prominent after elongation ceases. Secondary xylem is of greater radial width in the stem sectors which include protoxylem. During early secondary growth, vessels develop in the secondary xylem only in these sectors. Nucleate fibers and rays constitute the remainder of the secondary xylem. The rays exhibit an organization noted in other plants of reduced growth habit. Some of these interpretations do not agree with those described for tomato in earlier studies, and they are discussed in relation to pertinent aspects of development.     

Compressive/Tensile Stresses and Lignified Cells as Resistance Components in Joints between Cladodes of Opuntia laevis (Cactaceae)

The Cactaceae are a diverse group of plants with a wide variety of morphologies. Many species of Opuntia have segmented stems in which terminal cladodes may be separated from main-stem cladodes with varying amounts of resistance. From a geometric approach, derivations were used to calculate normal (axial and bending) and shear (transverse force and torque) stresses at joints due to the weight of the cladodes. Normal and shear stresses act perpendicular and parallel to (along) the cross sections of joints, respectively. Normal stress caused by bending was >10 times that of the mean value of any other stress. Analyses were performed to determine the relationship between maximum normal stress and the amount of lignified xylem cells. Such cells had thicker cell walls compared with the various other cells of stem joints that had thin cell walls and that thus would provide the most resistance to normal stresses. An analogy was made between cactus joints and a composite beam with reinforcing rods. In such joints, thin-walled parenchyma cells might be analogous to concrete that has little resistance to tensile stress, while the thick-walled, lignified xylem cells would be analogous to reinforcing rods. There were statistically significant relationships between normal stresses (from bending and axial loads) and mean percentage of lignified xylem cells (r=0.73) and between normal stresses and total areas of lignified xylem cells (r=0.65) (more stress, more reinforcing xylem cells). Tensile portions of cactus joints had 23% lignified xylem cells, while compressive portions had only 10% lignified xylem cells in joint areas (more tension, more reinforcing xylem cells). In addition, tensile joint tissues had two to three times more thick-walled, lignified xylem cells in the outer 30% of the radius compared with other joint tissues types (more reinforcing near the surface). To our knowledge, this is the first publication to present mechanical stresses at stem joints of cacti and the first to relate these stresses to characteristics of resisting tissues in the joints of a cactus.     

Abscisic acid triggers whole-plant and fruit-specific mechanisms to increase fruit calcium uptake and prevent blossom end rot development in tomato fruit

Calcium (Ca) uptake into fruit and leaves is dependent on xylemic water movement, and hence presumably driven by transpiration and growth. High leaf transpiration is thought to restrict Ca movement to low-transpiring tomato fruit, which may increase fruit susceptibility to the Ca-deficiency disorder, blossom end rot (BER). The objective of this study was to analyse the effect of reduced leaf transpiration in abscisic acid (ABA)-treated plants on fruit and leaf Ca uptake and BER development. Tomato cultivars Ace 55 (Vf) and AB2 were grown in a greenhouse environment under Ca-deficit conditions and plants were treated weekly after pollination with water (control) or 500 mg l(-1) ABA. BER incidence was completely prevented in the ABA-treated plants and reached values of 30-45% in the water-treated controls. ABA-treated plants had higher stem water potential, lower leaf stomatal conductance, and lower whole-plant water loss than water-treated plants. ABA treatment increased total tissue and apoplastic water-soluble Ca concentrations in the fruit, and decreased Ca concentrations in leaves. In ABA-treated plants, fruit had a higher number of Safranin-O-stained xylem vessels at early stages of growth and development. ABA treatment reduced the phloem/xylem ratio of fruit sap uptake. The results indicate that ABA prevents BER development by increasing fruit Ca uptake, possibly by a combination of whole-plant and fruit-specific mechanisms.     

Stomatal factors and vulnerability of stem xylem to cavitation in poplars

The relationships between the vulnerability of stem xylem to cavitation, stomatal conductance, stomatal density, and leaf and stem water potential were examined in six hybrid poplar (P38P38, Walker, Okanese, Northwest, Assiniboine and Berlin) and balsam poplar (Populus balsamifera) clones. Stem xylem cavitation resistance was examined with the Cavitron technique in well-watered plants grown in the greenhouse. To investigate stomatal responses to drought, plants were subjected to drought stress by withholding watering for 5 (mild drought) and 7 (severe drought) days and to stress recovery by rewatering severely stressed plants for 30 min and 2 days. The clones varied in stomatal sensitivity to drought and vulnerability to stem xylem cavitation. P38P38 reduced stomatal conductance in response to mild stress while the balsam poplar clone maintained high leaf stomatal conductance under more severe drought stress conditions. Differences between the severely stressed clones were also observed in leaf water potentials with no or relatively small decreases in Assiniboine, P38P38, Okanese and Walker. Vulnerability to drought-induced stem xylem embolism revealed that balsam poplar and Northwest clones reached loss of conductivity at lower stem water potentials compared with the remaining clones. There was a strong link between stem xylem resistance to cavitation and stomatal responsiveness to drought stress in balsam poplar and P38P38. However, the differences in stomatal responsiveness to mild drought suggest that other drought-resistant strategies may also play a key role in some clones of poplars exposed to drought stress.     

Visualization of lateral water transport pathways in soybean by a time of flight-secondary ion mass spectrometry cryo-system

Water movement between cells in a plant body is the basic phenomenon of plant solute transport; however, it has not been well documented due to limitations in observational techniques. This paper reports a visualization technique to observe water movement among plant cells in different tissues using a time of flight-secondary ion mass spectrometry (Tof-SIMS) cryo-system. The specific purpose of this study is to examine the route of water supply from xylem to stem tissues. The maximum resolution of Tof-SIMS imaging was 1.8 μm (defined as the three pixel step length), which allowed detection of water movement at the cellular level. Deuterium-labelled water was found in xylem vessels in the stem 2.5 min after the uptake of labelled water by soybean plants. The water moved from the xylem to the phloem, cambium, and cortex tissues within 30-60 min after water absorption. Deuterium ion counts in the phloem complex were slightly higher than those in the cortex and cambium tissue seen in enlarged images of stem cell tissue during high transpiration. However, deuterium ion counts in the phloem were lower than those in the cambium at night with no evaporative demand. These results indicate that the stem tissues do not receive water directly from the xylem, but rather from the phloem, during high evaporative demand. In contrast, xylem water would be directly supplied to the growing sink during the night without evaporative demand.     

Xylem acoustic emissions and water relations of Calluna vulgaris L. at two climatological regions of Britain

The acoustic emission rate (as a result of xylem cavitation), stem water content, stomatal conductance, and leaf water potential of heather (Calluna vulgaris L. Hull) were compared over a period of 18 months at two contrasting sites in Britain. The sites were Thursely Common in south-east England, and Flanders Moss in central Scotland (average rainfall of 600 mm per year and 1500 mm per year, respectively). In the first year of study (1992) the natural difference between the sites was amplified by a severe drought affecting south-east England. The relative water content of the xylem (RWC) did not differ between the sites, and did not fluctuate seasonally, despite the drought at Thursely. Acoustic emissions were detected at both sites, but the incidence was low. The same range of foliage water potential and stomatal conductance was found at both sites. The leaf area to sapwood area ratio at the dry site was half that at the wet site, and it is suggested that this adjustment of the transpiring area relative to the water conducting area facilitates the close physiological state of the plants at the two sites. The possible occurrence of aerial refilling and freezing induced xylem cavitation is also discussed.     

Hygrochastic capsule dehiscence supports safe site strategies in New Zealand alpine Veronica (Plantaginaceae)

Background and Aims

Hygrochasy is a capsule-opening mechanism predominantly associated with plants in arid habitats, where it facilitates spatially and temporally restricted dispersal. Recently, hygrochastic capsules were described in detail for the first time in alpine Veronica in New Zealand. The aim of the present study was to investigate whether hygrochastic capsules are an adaptation of alpine Veronica to achieve directed dispersal to safe sites. We expect that by limiting dispersal to rainfall events, distances travelled by seeds are short and confine them to small habitat patches where both seedlings and adults have a greater chance of survival.

Methods

Dispersal distances of five hygrochastic Veronica were measured under laboratory and field conditions and the seed shadow was analysed. Habitat patch size of hygrochastic Veronica and related non-hygrochastic species were estimated and compared.

Key Results

Dispersal distances achieved by dispersal with raindrops did not exceed 1 m but weather conditions could influence the even distribution of seeds around the parent plant. Compared with related Veronica species, hygrochastic Veronica mostly grow in small, restricted habitat patches surrounded by distinctly different habitats. These habitat patches provide safe sites for seeds due to their microtopography and occurrence of adult cushion plants. Non-hygrochastic Veronica can be predominantly found in large habitats without clearly defined borders and can be spread over long distances along rivers.

Conclusions

The results suggest that hygrochasy is a very effective mechanism of restricting seed dispersal to rainfall events and ensuring short-distance dispersal within a small habitat patch. It appears that it is an adaptation for directed dispersal to safe sites that only exist within the parent habitat.     

The application of various anatomical techniques for studying the hydraulic network in tomato fruit pedicels

The abscission zone in fruit pedicels plays an important role in affecting not only water uptake in the developing fruit, but also in the transport of chemical signals from root to shoot. In order to characterize the hydraulic network of tomato fruit pedicels, we applied various techniques, including light, fluorescence microscopy, electron microscopy, maceration, tissue clearing, and X-ray computed tomography. Because of significant changes in xylem anatomy, the abscission zone in tomato fruit pedicels is illustrated to show a clear reduction in hydraulic conductance. Based on anatomical measurements, the theoretical axial xylem conductance was calculated via the Hagen–Poiseuille law, suggesting that the hydraulic resistance of the abscission zone increases at least two orders of magnitude compared to the pedicel zone near the stem. The advantages and shortcomings of the microscope techniques applied are discussed.     

胡杨木质部水分传导对盐胁迫的响应与适应北大核心CSCD

解析植物木质部导水率对逆境的响应和适应对促进植物抗逆性机理研究和受损植被恢复具有重要意义。该文以荒漠河岸林建群种胡杨(Populus euphratica)为研究对象,系统分析了胡杨幼株根、茎、叶水分传输通道对不同浓度盐胁迫的响应和适应。结果表明:(1)胡杨幼株根系对盐胁迫的敏感性高于茎和叶,盐胁迫下根系生长和根尖数显著受到抑制,根木质部易于发生栓塞,导水率明显降低。(2)胡杨幼株茎木质部导水率对盐胁迫的响应依盐浓度而定,轻度(0.05 mol·L–1 Na Cl)和中度(0.15 mol·L–1 Na Cl)盐胁迫下,胡杨可以通过协调导管输水的有效性和安全性来调节木质部的导水率,维持植物正常生长;重度(0.30 mol·L–1 Na Cl)盐胁迫下,胡杨茎木质部导管输水有效性和安全性均明显降低,木质部导水率显著下降,并伴随叶片气孔导度的显著降低,从而严重抑制了胡杨的光合和生长。     

Differential allocation of constitutive and induced chemical defenses in pine tree juveniles: a test of the optimal defense theory

Optimal defense theory (ODT) predicts that the within-plant quantitative allocation of defenses is not random, but driven by the potential relative contribution of particular plant tissues to overall fitness. These predictions have been poorly tested on long-lived woody plants. We explored the allocation of constitutive and methyl-jasmonate (MJ) inducible chemical defenses in six half-sib families of Pinus radiata juveniles. Specifically, we studied the quantitative allocation of resin and polyphenolics (the two major secondary chemicals in pine trees) to tissues with contrasting fitness value (stem phloem, stem xylem and needles) across three parts of the plants (basal, middle and apical upper part), using nitrogen concentration as a proxy of tissue value. Concentration of nitrogen in the phloem, xylem and needles was found to be greater higher up the plant. As predicted by the ODT, the same pattern was found for the concentration of non-volatile resin in the stem. However, in leaf tissues the concentrations of both resin and total phenolics were greater towards the base of the plant. Two weeks after MJ application, the concentrations of nitrogen in the phloem, resin in the stem and total phenolics in the needles increased by roughly 25% compared with the control plants, inducibility was similar across all plant parts, and families differed in the inducibility of resin compounds in the stem. In contrast, no significant changes were observed either for phenolics in the stems, or for resin in the needles after MJ application. Concentration of resin in the phloem was double that in the xylem and MJ-inducible, with inducibility being greater towards the base of the stem. In contrast, resin in the xylem was not MJ-inducible and increased in concentration higher up the plant. The pattern of inducibility by MJ-signaling in juvenile P. radiata is tissue, chemical-defense and plant-part specific, and is genetically variable.     

Gradients and dynamics of inner bark and needle osmotic potentials in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst)

Preconditions of phloem transport in conifers are relatively unknown. We studied the variation of needle and inner bark axial osmotic gradients and xylem water potential in Scots pine and Norway spruce by measuring needle and inner bark osmolality in saplings and mature trees over several periods within a growing season. The needle and inner bark osmolality was strongly related to xylem water potential in all studied trees. Sugar concentrations were measured in Scots pine, and they had similar dynamics to inner bark osmolality. The sucrose quantity remained fairly constant over time and position, whereas the other sugars exhibited a larger change with time and position. A small osmotic gradient existed from branch to stem base under pre‐dawn conditions, and the osmotic gradient between upper stem and stem base was close to zero. The turgor in branches was significantly driven by xylem water potential, and the turgor loss point in branches was relatively close to daily minimum needle water potentials typically reported for Scots pine. Our results imply that xylem water potential considerably impacts the turgor pressure gradient driving phloem transport and that gravitation has a relatively large role in phloem transport in the stems of mature Scots pine trees.     

中亚热带喀斯特常绿落叶阔叶混交林典型树种的木质部解剖与功能特征分析

树木木质部主要由导管、纤维和薄壁组织组成, 分别具有运输、支撑和贮存的生理功能。由于木质部空间限制, 一种组织比例的增加会导致其他组织比例的降低, 因而可能表现出权衡关系。分析木质部组织比例和权衡关系有助于了解植物的生理生态适应性。该研究选择中亚热带喀斯特常绿落叶阔叶混交林21种典型树种(10种落叶树种, 11种常绿树种), 测定枝条木质部各组织比例, 计算水力相关指标并分析性状之间的相关性。结果表明: (1)与全球木质部解剖数据对比分析, 喀斯特树种木质部趋向具有较高比例的薄壁组织; (2)喀斯特树种导管组织比例与薄壁和纤维组织比例之间没有显著的相关性, 但是薄壁和纤维组织比例之间有显著的权衡关系; (3)常绿和落叶树种的木质部水力运输安全性(导管壁加固系数)和效率性(理论导水率)均具有显著的权衡关系, 但是这两个类群线性回归的截距存在显著差异, 即在相同的理论导水率条件下, 落叶树种比常绿树种具有较高的导管壁加固系数(安全性), 可能与常绿树种具有更多的轴向薄壁组织有关。喀斯特树种木质部解剖特征表明薄壁组织的贮存功能对喀斯特树种(尤其是常绿树种)的干旱适应具有重要作用。     

Maize stomatal conductance in the field: its relationship with soil and plant water potentials, mechanical constraints and ABA concentration in the xylem sap

Abstract. Stomatal conductance, leaf water potential, soil water potential and concentration of abscisic acid (ABA) in the xylem sap were measured on maize plants growing in the field, in two treatments with contrasting soil structures. Soil compaction affected the stomatal conductance, but this effect was no longer observed if the soil water potential was increased by irrigation. Differences in leaf water potential did not account for the differences in conductance between treatments. Conversely, the relationship between stomatal conductance and concentration of ABA in the xylem sap was consistent during the experiment. The proposed interpretation is that stomatal conductance was controlled by the root water potential via an ABA message. Control of the stomatal conductance by the leaf water potential or by an effect of mechanical stress on the roots is unlikely.