Architecture and growth of an annual plant <Emphasis Type="Italic">Chenopodium album</Emphasis> in different light climates

Light climates strongly influence plant architecture and mass allocation. Using the metamer concept, we quantitatively described branching architecture and growth of Chenopodium album plants grown solitarily or in a dense stand. Metamer is a unit of plant construction that is composed of an internode and the upper node with a leaf and a subtended axillary bud. The number of metamers on the main-axis stem increased with plant growth, but did not differ between solitary and dense-stand plants. Solitary plants had shorter thicker internodes with branches larger in size and number than the plant in the dense stand. Leaf area on the main stem was not different. Larger leaf area in solitary plants was due to a larger number of leaves on branches. Leaf mass per area (LMA) was higher in solitary plants. It did not significantly differ between the main axis and branches in solitary plants, whereas in the dense stand it was smaller on branches. Dry mass was allocated most to leaves in solitary plants and to stems in the dense stand in vegetative growth. Reproductive allocation was not significantly different. Branch/main stem mass ratio was higher in solitary than dense-stand plants, and leaf/stem mass ratio higher in branches than in the main axis. Nitrogen use efficiency (NUE) (dry mass growth per unit N uptake) was higher and light use efficiency (LUE) (dry mass growth per unit light interception) was lower in the plant grown solitarily than in the dense stand.     

The effects of mechanical stress and spectral shading on the growth and allocation of ten genotypes of a stoloniferous plant

BACKGROUND AND AIMS: Because plants protect each other from wind, stand density affects both the light climate and the amount of mechanical stress experienced by plants. But the potential interactive effects of mechanical stress and canopy shading on plant growth have rarely been investigated and never in stoloniferous plants which, due to their creeping growth form, can be expected to respond differently to these factors than erect plants. METHODS: Plants of ten genotypes of the stoloniferous species Potentilla reptans were subjected to two levels of mechanical stress (0 or 40 daily flexures) and two levels of spectral shading (15 % of daylight with a red:far red ratio of 0.3 vs. 50 % daylight and a red:far red ratio of 1.2). KEY RESULTS: Mechanically stressed plants produced more leaves with shorter more flexible petioles, more roots, and more but less massive stolons. Responses to spectral shading were mostly in the opposite direction to thigmomorphogenesis, including the production of thinner, taller petioles made of more rigid tissue. The degree of thigmomorphogenesis was either independent of light climate or stimulated by spectral shading. At the genotypic level there were no clear correlations between responses to shade and mechanical stress. CONCLUSIONS: These results suggest that in stoloniferous plants mechanical stress results in clones with a more compact, shorter shoot structure and more roots. This response does not appear to be suppressed by canopy shading, which suggests that wind shielding (reduced mechanical stress) by neighbours in dense vegetation serves as a cue that induces shade avoidance responses such as increased petiole elongation.     

Thigmomorphogenesis: Field and laboratory studies of Abies fraseri in response to wind or mechanical perturbation

Field- and greenhouse-grown Abies fraseri (Pursh) Poir. (Fraser fir) were analyzed for wind- or mechanically-induced flexure changes. These changes included inhibition of stem and needle elongation, reinforcement of branch bases around the stem, and increased radial growth in the direction of the mechanical perturbation (MP). Mature trees exposed to high wind conditions were severely flag-formed. These modified tree crowns had a lower drag than crowns of non-flag formed trees in wind-tunnel tests. In both field-grown and greenhouse-grown A. fraseri , MP induced a decrease in flexibility and increased elasticity of the stems. The increased radial growth of the stems overrode the increase in elasticity, resulting in the overall decrease in flexibility. The increase in radial growth caused by wind or mechanical flexure was due to greater cell divisions of the vascular cambium, resulting in increased numbers of tracheids. The decrease in stem elongation in these trees was due, at least in part, to a decrease in tracheid length. The potential biological and mechanical significance of these induced growth changes in trees are addressed. The data support the thigmomorphogenetic theory, which states that plants respond to wind and other mechanical perturbations in a way that is favorable to the plant for continued survival in windy environments.     

Stem extension and mechanical stability of Xanthium canadense grown in an open or in a dense stand

Background and Aims

Plants in open, uncrowded habitats typically have relatively short stems with many branches, whereas plants in crowded habitats grow taller and more slender at the expense of mechanical stability. There seems to be a trade-off between height growth and mechanical stability, and this study addresses how stand density influences stem extension and consequently plant safety margins against mechanical failure.

Methods

Xanthium canadense plants were grown either solitarily (S-plants) or in a dense stand (D-plants) until flowering. Internode dimensions and mechanical properties were measured at the metamer level, and the critical buckling height beyond which the plant elastically buckles under its own weight and the maximum lateral wind force the plant can withstand were calculated.

Key Results

Internodes were longer in D- than S-plants, but basal diameter did not differ significantly. Relative growth rates of internode length and diameter were negatively correlated to the volumetric solid fraction of the internode. Internode dry mass density was higher in S- than D-plants. Young''s modulus of elasticity and the breaking stress were higher in lower metamers, and in D- than in S-plants. Within a stand, however, both moduli were positively related to dry mass density. The buckling safety factor, a ratio of critical buckling height to actual height, was higher in S- than in D-plants. D-plants were found to be approaching the limiting value 1. Lateral wind force resistance was higher in S- than in D-plants, and increased with growth in S-plants.

Conclusions

Critical buckling height increased with height growth due mainly to an increase in stem stiffness and diameter and a reduction in crown/stem mass ratio. Lateral wind force resistance was enhanced due to increased tissue strength and diameter. The increase in tissue stiffness and strength with height growth plays a crucial role in maintaining a safety margin against mechanical failure in herbaceous species that lack the capacity for secondary growth.     

Not only light quality but also mechanical stimuli are involved in height convergence in crowded Chenopodium album stands

? In crowded stands, height is often similar among dominant plants, as plants adjust their height to that of their neighbours (height convergence). We investigated which of the factors, light quality, light quantity and mechanical stimuli, is primarily responsible for stem elongation and height convergence in crowded stands. ? We established stands of potted Chenopodium album plants. In one stand, target plants were surrounded by artificial plants that were painted black to ensure that the light quality was not modified by their neighbours. In a second stand, target plants were surrounded by real plants. In both stands, one-half of the target plants were anchored to stakes to prevent flexing by wind. The target plants were lifted or lowered by 10?cm to test whether height convergence was affected by the different treatments. ? Stem length was affected by being surrounded by artificial plants, anchoring and pot elevation, indicating that light quality, light quantity and mechanical stimuli all influenced stem elongation. Height convergence did not occur in the stand with artificial plants or in anchored plants. ? We conclude that light quality and mechanical stimuli are important factors for the regulation of stem growth and height convergence in crowded stands.     

Root biomechanics and whole-plant allocation patterns: responses of tomato plants to stem flexure

Cherry tomato plants (Lycopersicon esculentum Mill.) were grownwith or without stem flexure similar to that caused by windin order to determine whether stem flexure affects whole-plantbiomass allocation and increases the ability of a plant to withstandwind- induced forces. After 6 weeks of flexing (1 mm, 6 days/week),whole plants were harvested. The main differences found betweentreatments were in the primary shoot/root axis. The stem wassignificantly shorter and wider near the shoot/root junctionin flexed than control plants, both above- and below- ground.Flexed plants had significantly higher root/shoot dry weightratios than controls, but flexed plants and controls did notdiffer significantly in total leaf area, root length, or totalbiomass. Lateral roots from the top 2 cm of the taproot werenot affected by the flexing treatment for any of the factorsstudied: number of laterals, proximal diameter, elastic modulus,stress at failure, or work to failure. Lastly, the force requiredto uproot flexed plants did not differ significantly from thatfor controls. However, because their stems were shorter, flexedplants would have been subjected to smaller stem bending momentsand thus less stress near their root crowns than would controls.Moreover, flexed plants have wider stem bases, and should thusbe better able to resist the forces that affect stems. Thissuggests that in a windy situation, plants that have previouslybeen subjected to flexing could potentially withstand more forcethan unflexed controls. Key words: Anchorage, root, wind, mechanical stimulation, tomato     

Thigmomorphogenesis: Anatomical, morphological and mechanical analysis of genetically different sibs of Pinus taeda in response to mechanical perturbation

Twenty-three open pollinated families (half-sibs) and four controlled pollinated families (full-sibs) of Pinus taeda L. (loblolly pine) were grown in a greenhouse and analyzed for changes induced by mechanical perturbation (MP). These changes included inhibition of stem and needle elongation, bracing of branch nodes, and increased radial growth in the direction of the MP. Inhibition of stem elongation was the least variable feature measured. Leaf extension and stem diameter were highly variable between half-sibs. MP induced increased drag in greenhouse grown P. taeda in wind-tunnel tests. In P. taeda , MP induced decreased flexibility and increased elasticity and plasticity of the stem. The increased radial growth of the stems overrode the increase in elasticity, resulting in an overall decrease in flexibility. MP trees had a higher rupture point than non-MP controls. Increased radial growth is a result of more rapid cell divisions of the vascular cambium, resulting in increased numbers of tracheids. The decreased leader growth is partly due to a decreased tracheid length in response to MP.     

Growth and nitrogen use in Xanthium canadense grown in an open or in a dense stand

Plants develop branches profusely when grown solitarily, while less so when grown in a dense stand. Such changes in architecture are associated with changes in dry mass allocation and nitrogen use. Here, we studied what traits in plant growth and nitrogen use were influenced by different light climates in the stand. Annual plants (Xanthium canadense) were grown solitarily or in a dense stand. Dry mass growth was analyzed as the product of the net assimilation rate (NAR) and leaf area (LA). Nitrogen use efficiency (NUE) was analyzed as the product of nitrogen productivity (NP) and the mean residence time (MRT) of nitrogen. These growth variables were further factorized into their components. Solitary plants maintained a high NAR, whereas plants in the dense stand decreased the NAR due to mutual shading. Plants in the dense stand developed a larger LA with a higher specific leaf area than solitary plants. Solitary plants had higher NUE due to higher NP. A temporal increase in NUE was attributed to the increase in MRT of nitrogen. Light climate was different between solitary and dense-stand plants, but they took up a comparable amount of nitrogen and used it differently in response to the given light climate. NUE was thus demonstrated to be a useful tool for analyzing the mechanism leading to different N use in plant growth.     

Traits of Heracleum sosnowskyi Plants in Monostand on Invaded Area

The ability of giant hogweeds to form monodominant communities and even pure monostands in invaded areas has been well documented. Understanding of the mechanisms leading to monostand formation can aid in determining the limitations of existing community ecology models and establishing an effective management plan for invasive species elimination. The aim of this observational study was to investigate traits of Heracleum sosnowskyi plants (demography, canopy structure, morphology and physiology) of the plants in a pure stand in an invaded area useful for understanding potential monostand formation mechanisms. All measurements were performed in one typical Heracleum sosnowskyi monostand located in an abandoned agriculture field located in Syktyvkar city suburb (North-east Russia). This monostand consisted of five main plant growth stages: seed, seedling, juvenile, vegetative adult, and generative adult. Plants of all stages began to grow simultaneously shortly after the snowmelt, at the same time as spring ephemeral plant species grew. The density of generative plants did not change during the vegetation period, but the density of the other plant stages rapidly decreased after the formation of a tall (up to 2–2.5 m) and dense (Leaf area index up to 6.5) canopy. The canopy captured approximately 97% of the light. H. sosnowskyi showed high (several orders of magnitude higher than average taiga zone grasses) photosynthetic water use efficiency (6–7 μM CO₂/μM H₂O). Formation of H. sosnowskyi monostands occurs primarily in disturbed areas with relatively rich and well-moistened soils. Early commencement of growth, rapid formation of a dense canopy, high efficiency of light and water use during photosynthesis, ability of young plants to survive in low light conditions, rapid recovery of above-ground plant parts after damage, and the high density of the soil seed bank are the most important traits of H. sosnowskyi plants for monostand formation in invaded areas.     

Phenotypic plasticity of stem elongation in two ecotypes of Stellaria longipes: the role of ethylene and response to wind

Using two ecotypes of Stellaria longipes an alpine form with low plasticity and a prairie form with high plasticity, we investigated whether ethylene was involved in the response to wind stress and might be important in controlling plasticity of stem elongation. Stem growth inhibition was positively correlated with concentration of ethephon application and elevation in ambient ethylene in alpine ecotypes, whereas stem growth in prairie plants was stimulated by low ethephon concentrations. When treated with high AVG, the effects were reversed: alpine plant growth was promoted and prairie plant growth was inhibited. Prairie plants exhibited a daily rhythm in ethylene evolution which increased and peaked at 1500 h, and which was absent in alpine plants. Ethylene evolution did not change significantly during the first 2 weeks of growth in alpine plants, whereas ethylene in prairie plants increased significantly during periods of rapid stem elongation. Wind treatment inhibited growth in both ecotypes, but only alpine plants showed a recovery of growth to control levels when wind stressed plants were pretreated with STS. In addition, only alpine plants showed an increase in ethylene evolution in response to wind simulation, whereas prairie plant ethylene evolution did not deviate from rhythms observed in unstressed plants. We concluded that ethylene dwarfs stems in alpine S. longipes in response to wind stress. However, low levels of ethylene may stimulate growth in prairie ecotypes and act independently of wind stress intensity. The contrasting ability to synthesize and respond to ethylene can account for part of the difference in plasticity documented between the two ecotypes.     

Interactive effects of lateral shade and wind on stem allometry, biomass allocation, and mechanical stability in Abutilon theophrasti (Malvaceae)

The effects of lateral shade and wind on stem allometry, whole-plant biomass allocation, and mechanical stability were examined for Abutilon theophrasti in a fully factorial glasshouse experiment. Lateral shade from neighboring plants increased stem height by 33% relative to control plants grown individually, despite a decrease in plant dry mass. Intermittent wind decreased stem height by 18% in unshaded plants, but by only 3% in shaded plants. Surprisingly, both lateral shade and wind caused decreases in stem diameter, even with diameter controlled for height, resulting in low diameter?:?height ratios in wind-treated plants relative to untreated plants. Under shade, wind-treated plants had higher root allocation than untreated plants, which allowed wind-treated shade plants to compensate for a low diameter?:?height ratio. This did not occur in the absence of shade, where stem tissue density and root allocation of wind-treated plants did not exceed that of untreated plants. Nevertheless, wind-treated plants experienced low drag relative to untreated plants due to a lower leaf area. Consequently, stem deflections of wind-treated plants did not exceed those of untreated plants at any given windspeed. Our results document a complex interaction between shade and wind on plant morphology and suggest that the nature of this interaction is generally that lateral shade acts to reduce or eliminate thigmomorphogenic responses.     

Posture control and skeletal mechanical acclimation in terrestrial plants: implications for mechanical modeling of plant architecture

Self-supporting plant stems are slender, erect structures that remain standing while growing in highly variable mechanical environments. Such ability is not merely related to an adapted mechanical design in terms of material-specific stiffness and stem tapering. As many terrestrial standing animals do, plant stems regulate posture through active and coordinated control of motor systems and acclimate their skeletal growth to prevailing loads. This analogy probably results from mechanical challenges on standing organisms in an aerial environment with low buoyancy and high turbulence. But the continuous growth of plants submits them to a greater challenge. In response to these challenges, land plants implemented mixed skeletal and motor functions in the same anatomical elements. There are two types of kinematic design: (1) plants with localized active movement (arthrophytes) and (2) plants with continuously distributed active movements (contortionists). The control of these active supporting systems involves gravi- and mechanoperception, but little is known about their coordination at the whole plant level. This more active view of the control of plant growth and form has been insufficiently considered in the modeling of plant architecture. Progress in our understanding of plant posture and mechanical acclimation will require new biomechanical models of plant architectural development.     

Ethylene sensitivity affects changes in growth patterns, but not stem properties, in response to mechanical stress in tobacco

Plant responses to mechanical stress (e.g. wind or touch) involve a suite of physiologic and developmental changes, collectively known as thigmomorphogenesis, including reductions in height increment, Young's modulus of stems, shoot growth, and seed production, and increased stem girth and root growth. A role of the phytohormone ethylene in thigmomorphogenesis has been proposed but the extent of this involvement is not entirely clear. To address this issue, wild-type (WT) and ethylene-insensitive transgenic (Tetr) tobacco ( Nicotianum tabacum ) plants were subjected to three levels of mechanical stress: 0, 25 and 75 daily flexures. Flexed plants produced shorter, thicker stems with a lower Young's modulus than non-flexed ones, and these responses occurred independently of genotype. This suggests that ethylene does not play a role in thigmomorphogenesis-related changes in stem characteristics in tobacco. The effect of mechanical stress on dry mass increment (growth), on the other hand, differed between the genotypes: in the WT plants, shoot growth but not root growth was reduced under mechanical stress, resulting in reduced total growth and increased root mass fractions. In the Tetr plants, neither shoot nor root growth were affected. This suggests that ethylene is involved in the inhibition of tobacco shoot growth under mechanical stress.     

The effects of mechanical stress on the growth, differentiation, and paracrine factor production of cardiac stem cells

Stem cell therapies have been clinically employed to repair the injured heart, and cardiac stem cells are thought to be one of the most potent stem cell candidates. The beating heart is characterized by dynamic mechanical stresses, which may have a significant impact on stem cell therapy. The purpose of this study is to investigate how mechanical stress affects the growth and differentiation of cardiac stem cells and their release of paracrine factors. In this study, human cardiac stem cells were seeded in a silicon chamber and mechanical stress was then induced by cyclic stretch stimulation (60 cycles/min with 120% elongation). Cells grown in non-stretched silicon chambers were used as controls. Our result revealed that mechanical stretching significantly reduced the total number of surviving cells, decreased Ki-67-positive cells, and increased TUNEL-positive cells in the stretched group 24 hrs after stretching, as compared to the control group. Interestingly, mechanical stretching significantly increased the release of the inflammatory cytokines IL-6 and IL-1β as well as the angiogenic growth factors VEGF and bFGF from the cells in 12 hrs. Furthermore, mechanical stretching significantly reduced the percentage of c-kit-positive stem cells, but increased the expressions of cardiac troponin-I and smooth muscle actin in cells 3 days after stretching. Using a traditional stretching model, we demonstrated that mechanical stress suppressed the growth and proliferation of cardiac stem cells, enhanced their release of inflammatory cytokines and angiogenic factors, and improved their myogenic differentiation. The development of this in vitro approach may help elucidate the complex mechanisms of stem cell therapy for heart failure.     

Biomechanical study of the effect of a controlled bending on tomato stem elongation: global mechanical analysis

An experiment was designed to apply a controlled bending to a tomato stem and simultaneously to measure its effect on stem elongation. Stem elongation was measured over 2 d until steady and equal rates were obtained for the control and the treated plants. Thereafter, the basal part of the stem was submitted to a transient controlled bending at constant displacement rate using a motorized dynamometer. After load removal, stem elongation was again measured for 2 d. The tested plants were mature (height visible internodes) and only the basal part of the stem, which had already finished elongation, was loaded (hypocotyl and the first three internodes). A few minutes after the application of bending, elongation stopped completely for 60 min. Thereafter it took 120-1000 min to recover a rate of elongation similar to the control. The growth response was exclusively due to the bending of the basal part of the stem. It was shown that the side mechanical perturbations on the roots and on the stem tissues interacting directly with the clamp were not significantly involved on the elongation response. These results give evidence for mechanical perception and plant signalling from the basal stem to the upper elongating zone. However, none of the variables characterizing the global mechanical state of the bent part of the stem (i.e. the maximal force, bending moment, inclination, mean curvature of the stem, stored mechanical energy) could quantitatively explain the variability of the growth response. A more local mechanical analysis is therefore needed to elucidate how the mechanical stimulus is perceived.     

Environmental control of growth and development of Scrophularia marilandica

Summary Seedlings of Scrophularia marilandica were grown at different combinations of day/night temperature and photoperiod under controlled conditions. The species flowered in long days. The stems of plants grown at low temperature and short photoperiod failed to elongate. Treatment with gibberellic acid (GA₃) simulated the effect of increasing temperature and photoperiod and caused stem elongation in plants which would otherwise not have elongated. Application of GA₃ to plants grown at high temperature and long photoperiod resulted in increased stem elongation and flowering. The growth retardant (2-chloroethyl)trimethylammonium chloride (CCC) had little effect on rosette plants grown at low temperature and short photoperiod. Application of CCC to +GA₃ plants grown at a higher temperature and long photoperiod gave a significant increase in stem height. The interaction between temperature and applied GA is described in an experiment using plants grown at high and low temperatures for varying periods of time.This work was supported by National Science Foundation Grant GB 17483.     

Field measurements of wind speed and reconfiguration in Arundo donax (Poaceae) with estimates of drag forces

The giant reed (Arundo donax) is well known as a species that can withstand high wind loads without mechanical damage. To examine wind impact, profiles of vertical wind speeds in the plant's natural habitat (southern France) were measured at the edge and within a stand in the main wind direction. Wind speed was recorded simultaneously at five heights. For 75 measurements of within-canopy wind speed profiles, the attenuation coefficient was 4.4 ± 0.5, a value typical for plant stands with very dense canopies. Video recordings proved that A. donax becomes streamlined with increasing wind speed, reducing the projected surface area of leaves and stem. The total projected surface area is a function of wind speed and can be characterized by a second-order polynomial regression curve. For small wind velocities up to 1 m/s, the calculated drag force is proportional to the square of the wind speed. However, when A. donax plants are subjected to higher wind speeds (1.5-10 m/s), the drag force becomes directly proportional to the wind speed. Streamlining is a potentially important adaptation for withstanding high wind loads, especially for individual plants and plants at the edge of stands, whereas in dense stands streamlining probably plays a minor role.     

Thigmomorphogenesis: On the mechanical properties of mechanically perturbed bean plants

The mechanical properties of control and mechanically perturbed (MP) bean stems ( Phaseolus vulgaris L., ev. Cherokee wax) were compared. The rubbed plants were greatly hardened against mechanical rupture by previous MP. This hardening was due to a dramatic increase in the flexibility of the stems, but not in their stiffness. The MP-plants were able lo bend more than 90° without breaking, whereas the control plants broke after just slight bending. A comparison with other work reveals that different species utilize different tactics for achieving similar resistance to rupture due to mechanical stress.     

Development and Growth Form of the Neotropical Liana <Emphasis Type="Italic">Croton nuntians</Emphasis>: The Effect of Light and Mode of Attachment on the Biomechanics of the Stem

The neotropical liana Croton nuntians (Euphorbiaceae) can occur in a variety of different growth habits. Juvenile freestanding plants are mechanically stable without support and resemble morphologically young trees or shrubs, whereas adult plants are climbers. Ontogenetic variation of bending and torsion properties of different growth phases are analyzed by measurements of flexural stiffness, structural bending modulus, torsional stiffness and structural torsional modulus. Mechanical and anatomical data show two fundamentally different patterns for juvenile freestanding and adult climbing plants. In freestanding plants, mechanical properties and the contribution of cortex, wood, and pith to the stem cross-section vary only little during ontogeny as is typical for semi-self-supporting plants. In contrast, climbing plants become significantly more flexible during ontogeny, as is characteristic for lianas. This is accompanied by a transition to the formation of a less dense wood type with large diameter vessels and an increasing contribution of flexible tissues (less dense wood and cortex) to the cross-sectional area and the axial second moment of area of the stems. Depending on the environmental conditions, freestanding plants can differ considerably in their appearance due to differences in branching system or stem taper. Therefore the influence of light quantity, measured as percentage of canopy opening, on the mechanical properties and the stem anatomy was tested. Freestanding plants grown with strong shade are significantly more stiff in bending compared with plants grown with a moderate light environment.     

Mechanical Differences Between Free-standing and Supported Wheat Plants, Triticum aestivum L.

The effect of wind sway on the mechanical characteristics ofthe anchorage roots and the stem was investigated in maturewinter wheat (Triticum aestivumL., cv. Hereward). Wheat plantswere field-grown, either supported by a frame, which preventedwind sway, or unsupported (free-standing) and the morphologyand mechanical properties of the stems and the anchorage, ‘coronal’, roots were measured. Wind sway had little influence on either the stem height orear weight of the plants but did affect the mechanical propertiesof the stem. Stems of supported plants were weaker and moreflexible than the stems of free-standing plants. There werealso differences in the anchorage systems between the treatments:supported plants had just under half as many ‘coronal’ anchorage roots as the free-standing plants. This reducedthe anchorage strength of supported plants by a third. These differences in mechanical structure meant that the free-standingplants were more resistant to stem buckling and more resistantto anchorage failure. However, considering the difference inthe need for mechanical strength in plants from the two regimes,these differences were small. This suggests that wheat has inherentmechanical integrity and, as a monocotyledon with no secondarythickening, it differs little structurally between environments. Triticum aestivumL.; thigmomorphogenesis; anchorage; safety factor; mechanical stimulation