Control of growth of juvenile leaves of Eucalyptus globulus: effects of leaf age

Biophysical variables influencing the expansion of plant cells (yield threshold, cell wall extensibility and turgor) were measured in individual Eucalyptus globulus leaves from the time of emergence until cessation of growth. Leaf water relations variables and growth rates were determined as relative humidity was changed on an hourly basis. Yield threshold and cell wall extensibility were estimated from plots of leaf growth rate versus turgor. Cell wall extensibility was also measured by the Instron technique, and yield threshold was determined experimentally both by stress relaxation in a psychrometer chamber and by incubation in a range of polyethylene glycol solutions. Once emerging leaves reached approximately 5 cm(2) in size, increases in leaf area were rapid throughout the expansive phase and varied little between light and dark periods. Both leaf growth rate and turgor were sensitive to changes in humidity, and in the longer term, both yield threshold and cell wall extensibility changed as the leaf aged. Rapidly expanding leaves had a very low yield threshold and high cell wall extensibility, whereas mature leaves had low cell wall extensibility. Yield threshold increased with leaf age.     

Diurnal variation in leaf extension of Salix viminalis at two nitrogen supply rates

To investigate how nitrogen supply might affect the biophysical factors controlling diurnal variation in leaf extension, pot-grown Salix viminalis L. were supplied with nitrogen at a low relative addition rate of 0.05 g N g(-1) N day(-1) (low N) or were given free access to all nutrients (high N). Leaf extension, turgor pressure, turgor after stress relaxation and the plastic extensibility of leaf tissue were determined for growing leaves every 4 h during two days of clear skies in August. Plants in the high-N treatment had a significantly higher relative growth rate, dry weight, shoot/root ratio, leaf nitrogen concentration, total leaf area, final area of single leaves and epidermal cell size than plants in the low-N treatment. The periodicity of leaf extension was similar in both treatments with high values during the afternoon and early evening, and negligible values during the night and in the early morning. The maximum rate of leaf extension was higher in high-N than in low-N plants. Leaf water potential and leaf osmotic potential decreased in the morning and increased in the afternoon with highest values during the night. Calculated values of turgor pressure showed no consistent diurnal trend and did not correlate with the rate of leaf extension. There was no consistent difference in turgor between treatments. Turgor after stress relaxation varied diurnally. The difference between turgors before and after stress relaxation also varied diurnally and was largely in phase with the diurnal pattern of leaf extension. These data are consistent with either a causal role for growth turgor (difference between turgors before and after stress relaxation) in the regulation of cell expansion, or a diurnal variation in turgors after relaxation, attributable to different capacities for cell wall loosening at different times of day. Plastic extensibility of leaf tissue showed no diurnal pattern but consistently higher values were found in high-N than in low-N plants. We conclude that the effects of nitrogen supply on leaf water relations did not limit leaf extension, but that nitrogen supply did affect processes associated with cell wall loosening and enlargement. Nitrogen supply did not affect final values of turgor after relaxation, but it presumably affected the rate at which relaxation proceeded.     

Diurnal variation in extension growth of leaves of Salix viminalis

Diurnal variation in leaf extension and biophysical parameters of leaf growth were measured in young leaves from a stand of Salix viminalis L. in southern Sweden over a two-day period of clear skies during late July. Leaf growth rate (irreversible extension) was greatest during the late afternoon and early evening, falling to negligible values during the night and early morning.Leaf water potential and leaf osmotic potential showed declining values in the morning with subsequent recovery in the late afternoon. Diurnal variation in osmotic potential (-1.3 to -1.7 MPa) was small compared with that of leaf water potential (-0.1 to -1.2 MPa). Calculated values of leaf turgor pressure during the night (1.2 MPa) were double the midday values. Growth rate correlated poorly with turgor, which (except on one occasion) was always above a calculated value of yield turgor at 0.53 MPa.Diurnal variation in extension growth rate was large compared with that in plastic extensibility of leaf tissue as measured by an Instron technique. Values of extensibility were low and showed little diurnal variation, which is consistent with a proposed negative feedback of expansive growth rate on extensibility. Extension growth rate correlated well with air temperature, suggesting that the rate of leaf expansion may have been limited by a temperature-dependent rate of cell wall loosening.     

Biophysical constraints on leaf expansion in a tall conifer

The physiological mechanisms responsible for reduced extension growth as trees increase in height remain elusive. We evaluated biophysical constraints on leaf expansion in old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Needle elongation rates, plastic and elastic extensibility, bulk leaf water (Psi(L)) and osmotic (Psi(pi)) potential, bulk tissue yield threshold and final needle length were characterized along a height gradient in crowns of > 50-m-tall trees during the period between bud break and full expansion (May to June). Although needle length decreased with increasing height, there was no height-related trend in leaf plastic extensibility, which was highest immediately after bud break (2.9%) and declined rapidly to a stable minimum value (0.3%) over a 3-week period during which leaf expansion was completed. There was a significant positive linear relationship between needle elongation rates and plastic extensibility. Yield thresholds were consistently lower at the upper and middle crown sampling heights. The mean yield threshold across all sampling heights was 0.12 +/- 0.03 MPa on June 8, rising to 0.34 +/- 0.03 MPa on June 15 and 0.45 +/- 0.05 MPa on June 24. Bulk leaf Psi(pi) decreased linearly with increasing height at a rate of 0.004 MPa m(-1) during the period of most rapid needle elongation, but the vertical osmotic gradient was not sufficient to fully compensate for the 0.015 MPa m(-1) vertical gradient in Psi(L), implying that bulk leaf turgor declined at a rate of about 0.011 MPa m(-1) increase in height. Although height-dependent reductions in turgor appeared to constrain leaf expansion, it is possible that the impact of reduced turgor was mitigated by delayed phenological development with increasing height, which resulted in an increase with height in the temperature during leaf expansion.     

Changes in fruit sugar concentrations in response to assimilate supply,metabolism and dilution: a modeling approach applied to peach fruit (Prunus persica)

The influence of assimilate supply, metabolism and dilution on sugar concentrations in the mesocarp of peach (Prunus persica (L.) Batsch) fruit during the main stage of fruit enlargement was analyzed with the SUGAR model of Génard and Souty (1996). The model predicts the partitioning of carbon into sucrose, sorbitol, glucose and fructose in the mesocarp of peach fruit. Based on measured data and the model, we determined values for the relative rates of sugar transformation. These rates were constant, varied with time or varied with relative fruit growth rate, depending on the type of sugar. Equations were derived to describe these rates and incorporated into the SUGAR model. The model simulated the effects of changing assimilate supply and fruit volume on sugar concentrations. The set of equations for the SUGAR model was rewritten to include the three components influencing sugar concentrations: assimilate supply, metabolism and dilution. The sugar types differed in sensitivity to these components. Sucrose was highly sensitive to changes in assimilate supply and to the dilution effect; it was not subject to intense metabolic transformation. Sorbitol was the most important carbohydrate in fruit metabolism, which explains why the sorbitol concentration was always low despite the strong positive effect of assimilate supply. The reducing sugars constituted a transitory storage pool and their concentrations were closely related to metabolism.     

A carbon balance model of peach tree growth and development for studying the pruning response

We modeled tree responses to pruning on the basis of growth rules established on unpruned trees and a simple principle governing root-shoot interactions. The model, which integrates architectural and ecophysiological approaches, distinguishes four types of anatomical organs in a tree: rootstock, main axis, secondary axes and new roots. Tree structure is described by the position of secondary axes on the main axis. The main processes considered are plastochronal activity, branching, assimilate production, respiration and assimilate partitioning. Growth and development rules were based on measurements of two unpruned trees. The model was used to simulate growth of peach trees (Prunus persica (L.) Batsch) in their first growing season. Assuming that the equilibrium between roots and shoots tends to be restored after pruning, the response to removal of the main axis above the twentieth internode in mid-July was simulated and compared to the response measured in three pruned trees. The model fit the unpruned tree data reasonably well and predicted the main traits of tree behavior after pruning. Dry matter growth of the secondary axes of pruned trees was increased so that shoot seasonal carbon balance was hardly modified by pruning. Rhythmicity of growth was enhanced by pruning, and might result from variations induced in the root:shoot ratio. Variation in pruning severity had greater effects than variation in pruning date. A sensitivity analysis indicated that: (1) root-shoot partitioning was a critical process of the model; (2) tree growth was mainly dependent on assimilate availability; and (3) tree shape was highly dependent on the branching process.     

Leaf water relations of Eucalyptus globulus ssp. globulus and E. nitens: seasonal, drought and species effects

In August 1990, a 2-ha plantation was established in an area where rainfall (about 515 mm year(-1)) was insufficient to meet evaporative demand. On nine occasions between September 1991 and April 1993, pressure-volume curves were constructed for irrigated and rainfed Eucalyptus globulus ssp. globulus Labill. and E. nitens (Deane and Maiden) Maiden trees. During the experiment, rainfed trees experienced six periods when predawn water potential was significantly lower than that of irrigated trees. In early spring of 1991 and 1992, osmotic potentials at full turgor and turgor loss point in the irrigated E. nitens were significantly lower than at other times of the year, probably because of winter hardening. Water stress reduced osmotic potential and increased bulk elastic modulus in E. nitens, whereas the reverse occurred in E. globulus. However, treatment differences with respect to changes in osmotic and elastic properties were commonly overshadowed by interspecific differences. These were most apparent at the end of the sixth period of water stress when osmotic potentials at full and zero turgor were significantly higher and bulk elastic modulus and relative water content at turgor loss point were significantly lower in E. globulus than in E. nitens. We conclude that the drought-tolerance responses of E. globulus make it a more suitable species than E. nitens for establishment on sites where moderate water stress is experienced.     

Leaf water relations of Eucalyptus cloeziana and Eucalyptus argophloia in response to water deficit

Leaf water relations responses to limited water supply were determined in 7-month-old plants of a dry inland provenance of Eucalyptus argophloia Blakely and in a humid coastal provenance (Gympie) and a dry inland provenance (Hungry Hills) of Eucalyptus cloeziana F. Muell. Each provenance of E. cloeziana exhibited a lower relative water content at the turgor loss point, a lower apoplastic water content, a smaller ratio of dry mass to turgid mass and a lower bulk modulus of elasticity than the single provenance of E. argophloia. Osmotic potential at full turgor and water potential at the turgor loss point were significantly lower in E. argophloia and the inland provenance of E. cloeziana than in the coastal provenance of E. cloeziana. There was limited osmotic adjustment in response to soil drying in E. cloeziana, but not in E. argophloia. Between-species differences in water relations parameters were larger than those between the E. cloeziana provenances. Both E. cloeziana provenances maintained turgor under moderate water stress through a combination of osmotic and elastic adjustments. Eucalyptus argophloia had more rigid cell walls and reached lower water potentials with less reduction in relative water content than either of the E. cloeziana provenances, thereby enabling it to extract water from dryer soils.     

The supply of matrix containing glucomannans to the innermost surface of S2 layers is associated with changes in turgor pressure of differentiating tracheids in Cryptomeria japonica

We investigated the relationship between turgor pressure and diurnal differences in secondary wall formation of differentiating tracheids. Saplings of Cryptomeria japonica were grown in a growth chamber with 12-h light:12-h dark cycles, and the tangential strain on the inner bark surface was measured as an indicator of the volumetric changes of differentiating cells. The innermost surface of developing secondary walls was then observed using field emission scanning electron microscopy at 1-h intervals after both light and dark periods. Dramatic changes in the aspects of the innermost surface of developing secondary walls occurred 3h after the light was switched on and 4h after the light was switched off. The amorphous material containing glucomannans became evident when the differentiating cells became fully turgid during the dark period. Conversely, cellulose microfibrils became clearly visible when the cell volume was low during the light period. These results suggest that the diurnal periodicity in the supply of hemicellulose-containing matrix to developing secondary walls is associated with the changes in turgor pressure of differentiating tracheids that result from the change in light conditions during the photoperiodic cycle.     

针叶树木材细胞力学及纵向弹性模量计算——纵向弹性模量的理论模型

本文是关于针叶树木材弹性特性研究的理论分析。设定针叶树木材细胞主要由管胞和射线组成 ,根据管胞和射线细胞的解剖构造特征 ,建立了两端劈尖、矩形截面、中空的管胞模型与长方体状、中空的射线细胞模型。沿用MP层 (胞间层 初生壁 )内与 3S层 (次生壁的S1、S2、S3层 )内力学性质相同 ,而MP与 3S层间力学性质相异的假定 ,利用管胞和射线细胞模型 ,导出了管胞和射线细胞纵向弹性模量的计算公式。同时 ,根据管胞和射线细胞在针叶树木材中的排列规律 ,结合线形弹性体串并联的特性 ,进一步给出了计算针叶树木材试件宏观纵向弹性模量的方法。本项研究从木材细胞的结构和弹性特性出发 ,分析研究针叶树木材的宏观弹性行为 ,一方面可以获得关于针叶树木材弹性特性的认识 ,另一方面可以用于针叶树木材纵向弹性模量的计算和预测。     

Cellular basis for limitation of poplar leaf growth by water deficit

During the summers of 1986 and 1987, stem and leaf growth were measured on coppiced plants of Populus trichocarpa Torr. & A. Gray, P. deltoides Bartr. ex Marsh, and P. trichocarpa x deltoides growing in the field in Puyallup, WA. The trees were either irrigated periodically throughout the season, or grown without irrigation. In both treatments, stem volume at the end of the growing season was directly proportional to total leaf area in all three genotypes. The rate of individual leaf growth was reduced by lack of irrigation more in the parental species than in the hybrid. Only in the parental species did unirrigated trees have lower leaf water potentials (predawn and midday) than irrigated trees. However, stomatal conductances of all three genotypes were lower in unirrigated trees than in irrigated trees. Osmotic potentials of growing leaves of all three genotypes were also lower in unirrigated trees than in irrigated trees. As a consequence, turgor of growing leaves was as great in unirrigated trees as in irrigated trees, which indicates that turgor differences cannot explain the lower rates of leaf growth in the unirrigated trees. However, cell wall extensibility of leaves was lower in unirrigated trees than in irrigated trees, and the difference was greater in the parental species than in the hybrid. Unlike its effect on leaf area growth, irrigation increased stem volume growth of the hybrid and the parental species by a similar amount (12-16%).     

纳米压痕技术测量管胞次生壁S2层的纵向弹性模量和硬度

重点研究和详细介绍了利用纳米压痕技术测试管胞次生壁S2 层纵向弹性模量以及硬度的实验技术 ,并测试了人工林杉木早晚材管胞S2 层的纵向弹性模量和硬度。结果表明 :杉木晚材管胞S2 层的平均纵向硬度为0 390GPa ,弹性模量的平均值为 1 4 84 4GPa ;早材管胞S2 层的硬度和模量则小于晚材 ,平均值分别为 0 30 6GPa和9 82 3GPa。     

Influence of water-stress acclimation and <Emphasis Type="Italic">Tuber melanosporum</Emphasis> mycorrhization on <Emphasis Type="Italic">Quercus ilex</Emphasis> seedlings

Mycorrhizal and non-mycorrhizal holm oak (Quercus ilex L.) seedlings inoculated with black truffle (Tuber melanosporum) were grown under nursery conditions and subjected to drought hardening for 4 months in autumn and winter followed by irrigation for 10 days. Leaf water potential and stomatal conductance were monitored during the 4 months of drought. When the test was completed (March), measurements were made for each treatment (inoculated or non-inoculated), and watering regime (watered and water-stressed). Pressure–volume curves, osmotic potential at full turgor, osmotic potential at zero turgor and the tissue modulus of elasticity near full turgor were calculated. Mycorrhizal colonization and growth, and the content of the main mineral nutrients N, P, K, Ca and Mg were measured. Water stress affected plant growth, caused an elastic adjustment of the plant tissues, and decreased the P and K content, and inoculation improved the nitrogen content. Drought acclimation apparently achieved the goal of improving the drought tolerance of holm oak seedlings, without depressing ectomycorrhizal root colonization by T. melanosporum. José Antonio Rodríguez Barreal—Deceased     

Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. III. Diurnal patterns as influenced by vapor pressure deficit and internal water status

Pressure-volume curves were constructed and shoot water potentials measured for +20-year-old black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families growing on a moist site and a dry site at the Petawawa Research Forest, Ontario, to determine whether differences in diurnal water relations traits were related to productivity. To assess the basis for the observed diurnal patterns, we analyzed effects of environmental and internal water stress variables on diurnal water relations traits. Among the water relations traits examined, turgor pressure was the most sensitive, responding to site, family and environmental variables and displaying the strongest diurnal responses to varying soil water availability and atmospheric vapor pressure deficit (VPD). Overall, there was an 84% drop in turgor pressure with increasing VPD: turgor pressure fell 46% in response to the first 0.75 kPa increase in VPD, and 9.7% in response to a second 0.75 kPa increase in VPD. The families differed in water relations responses to moderate water stress, but not in responses to minor or more extreme water stresses. Thus, at a VPD of 0.5 kPa, there was an estimated 83% greater family difference in turgor pressure on the dry site compared with the moist site. Soil and atmospheric water stress appeared to exert effects in tandem to elicit these responses (r(2) = 0.728). A comparison of the mechanisms of response to water deficit indicated that osmotic adjustment was more important than change in cell wall elasticity. We used a conceptual water relations model to illustrate the differences between tolerant and intolerant families in their mechanisms of water stress response. We conclude that, because genetic responses to site factors are dynamic, the integrated response over time contributes to the observed genetic x environmental interaction in growth.     

基于纳米压痕技术的木材胶合界面力学行为

【目的】研究木材胶合界面的静态和动态力学行为,探讨树脂渗透对木材管胞壁层力学性能的影响,为木质复合材料制造工艺优化和增强改性提供理论依据。【方法】采用纳米压痕静态和动态力学测试技术(Nano-DMA),对针叶材火炬松与酚醛树脂(PF)、脲醛树脂(UF)胶黏剂所形成胶合界面区域各相材料的静态弹性模量、硬度、蠕变性能以及储能模量和损耗模量等力学行为进行分析。【结果】静态力学行为方面,在界面区域,PF和UF渗透进入管胞壁层后,木材管胞壁的弹性模量( E r)和硬度( H )提高;经PF渗透后,木材管胞壁的 E r和 H 分别增加7%和26%;Burgers蠕变力学模型可有效描述胶合界面区域管胞壁的纳米压痕蠕变特性,经树脂渗透后,木材管胞壁的瞬时弹性模量增加,黏弹性模量和黏性系数减小;在保载初期,PF界面区域木材管胞壁的蠕变柔量约下降60%,UF界面区域木材管胞壁的蠕变柔量约下降58%。动态力学行为方面,随着加载频率增加,界面材料的储能模量( E ′ r)逐渐增大,而损耗模量( E ″ r)和损耗因子(tan δ)呈减小趋势;当加载频率为10 Hz时,PF和UF树脂渗透使得管胞壁层的储能模量分别增加16%和29%。【结论】胶合界面区域胶黏剂进入管胞壁层,对木材管胞的静态力学性能具有增强作用,同时胶黏剂可提高管胞壁的短期抗蠕变能力;木材管胞壁具有较高的储能模量和损耗模量,而树脂的储能模量和损耗模量较低,经树脂渗透后,木材管胞壁的储能模量增加,但损耗模量和损耗因子呈下降趋势,可能对界面传递和分散应力产生不利影响。     

Growth, leaf morphology, water use and tissue water relations of Eucalyptus globulus clones in response to water deficit

Changes in leaf size, specific leaf area (SLA), transpiration and tissue water relations were studied in leaves of rooted cuttings of selected clones of Eucalyptus globulus Labill. subjected to well-watered or drought conditions in a greenhouse. Significant differences between clones were found in leaf expansion and transpiration. There was a significant clone x treatment interaction on SLA. Water stress significantly reduced osmotic potential at the turgor loss point (Pi0) and at full turgor (Pi100), and significantly increased relative water content at the turgor loss point and maximum bulk elastic modulus. Differences in tissue water relations between clones were significant only in the mild drought treatment. Among clones in the drought treatments, the highest leaf expansion and the highest increase in transpiration during the experiment were measured in those clones that showed an early and large decrease in Pi0 and Pi100.     

样品尺寸对竹材顺纹压缩力学性能的影响研究

以毛竹为研究对象,通过对不同尺寸的样品进行顺纹压缩测试,分析样品尺寸对竹材顺纹压缩模量和顺纹抗压强度的影响。结果显示：竹材的顺纹压缩模量与竹壁厚度、样品高度、弧度呈正相关关系,顺纹抗压强度与竹壁厚度、样品高度、弧度呈负相关关系。在弹性变形阶段,样品尺寸的增加会提高维管束的组织比量而增大竹材顺纹压缩模量的计算值,在塑性变形阶段,样品尺寸和弧度会导致提前失稳而降低竹材的顺纹抗压强度。     

Whole-plant adjustments in coconut (Cocos nucifera) in response to sink-source imbalance

Coconut (Cocos nucifera L.) is a perennial tropical monocotyledon that produces fruit continuously. The physiological function of the large amounts of sucrose stored in coconut stems is unknown. To test the hypothesis that reserve storage and mobilization enable the crop to adjust to variable sink-source relationships at the scale of the whole plant, we investigated the dynamics of dry matter production, yield and yield components, and concentrations of nonstructural carbohydrate reserves in a coconut plantation on Vanuatu Island in the South Pacific. Two treatments were implemented continuously over 29 months (April 2002 to August 2004): 50% leaf pruning (to reduce the source) and 100% fruit and inflorescence pruning (to reduce the sink). The pruning treatments had little effect on carbohydrate reserves because they affected only petioles, not the main reserve pool in the stem. Both pruning treatments greatly reduced dry matter production of the reproductive compartment, but vegetative growth and development were negligibly affected by treatment and season. Leaf pruning increased radiation-use efficiency (RUE) initially, and fruit pruning greatly reduced RUE throughout the experiment. Changes in RUE were negatively correlated with leaflet soluble sugar concentration, indicating feedback inhibition of photosynthesis. We conclude that vegetative development and growth of coconut show little phenotypic plasticity, assimilate demand for growth being largely independent of a fluctuating assimilate supply. The resulting sink-source imbalances were partly compensated for by transitory reserves and, more importantly, by variable RUE in the short term, and by adjustment of fruit load in the long term. Possible physiological mechanisms are discussed, as well as modeling concepts that may be applied to coconut and similar tree crops.     

Variation in surface conductance to water vapor diffusion in peach fruit and its effects on fruit growth assessed by a simulation model

Surface conductance to water vapor diffusion was measured in individual peach fruits (Prunus persica (L.) Batsch) and plotted as a function of fresh fruit mass for four cultivars. Surface conductance increased with fresh fruit mass, but the pattern differed with cultivar, and fruit-to-fruit variation occurred. Relationships between fruit mass and surface conductance were modeled by fitting mathematical equations to the data. The simulation model of Fishman and Génard (1998) was used to study dry mass and water components of fruit growth (1) when surface conductance varied with fruit size or was constant, and (2) when surface conductance values were high, moderate or low with respect to fruit mass. Increased surface conductance with fresh fruit mass resulted in fruit growth cessation. Fruits differing in surface conductance had similar dry mass. However, under well-watered conditions (stem water potential between -1 and -0.2 MPa), the water balance components of growth (osmotic and hydrostatic pressure, water potential and water balance) differed greatly and, as a result, the lower the surface conductance the greater the fresh fruit mass. These differences were buffered under drought conditions (stem water potential between -2.4 and -0.6 MPa).     

木塑复合材料性能影响因素分析

采用无损检测方法和静压试验,对由不同物料配比所得到的木塑复合材料动态弹性模量和静态弹性模量性能指标进行了检测,并分析了不同因素对材料弹性模量的影响程度,从而获知各因素对材料性能影响的主次顺序。由相关性分析得知,动态弹性模量与静态弹性模量之间具有很强的相关性。