Hisae Nagashima

Effects of Mechanical Stress and Plant Density on Mechanical Characteristics, Growth, and Lifetime Reproduction of Tobacco Plants

Plastic increases in stem elongation in dense vegetation are generally believed to be induced by canopy shading, but because plants protect each other from wind, shielding (reduced mechanical stress) could also play a role. To address this issue, tobacco Nicotiana tabacum plants were subjected to two levels of mechanical stress, 0 (control) or 40 (flexed) daily flexures, and grown solitarily, in a dense monostand (with plants of only one mechanical treatment), or in a mixed stand (flexed and control plants grown together). Flexed plants produced shorter and thicker stems with a lower Young’s modulus than control plants, while dense‐stand plants had relatively taller and thinner stems than solitary ones. Flexing effects on stem characteristics were independent of stand density. Growth, reproduction, and survival of solitary plants were not affected by flexing, while in the monostand growth was slightly reduced. But in the mixed stand, flexed plants were readily shaded by controls and had considerably lower growth, survival, and reproduction rates. These results suggest that wind shielding indeed plays a role in the plastic increase in stem elongation of plants in dense vegetation and that this response can have important consequences for competitive ability and lifetime seed production.

Leaf nitrogen distribution in relation to leaf age and photon flux density in dominant and subordinate plants in dense stands of a dicotyledonous herb

Abstract We studied the effects of photon flux density (PFD) and leaf position, a measure of developmental age, on the distribution of nitrogen content per unit leaf area (Narea) in plants of different heights, in dense stands grown at two nitrogen availabilities and in solitary plants of the erect dicotyledonous herb Xanthium canadense. Taller more dominant plants received higher PFD levels and experienced a larger difference in relative PFD between their youngest and oldest leaves than shorter subordinate plants in the stands. Differences in PFD between leaves of solitary plants were assumed to be minimal and differences in leaf traits, found for these plants, could thus be mainly attributed to an effect of leaf position. In the solitary plants, Narea decreased with leaf position while in the plants from the stands it decreased with decreasing relative PFD, indicating both factors to be important in determining the distribution of Narea. Due to the effect of leaf position on Narea, leaves of subordinate plants had a higher Narea than older leaves of dominant plants which were at the same height or slightly higher in the canopy. Consequently, the Narea distribution patterns of individual plants plotted as a function of relative PFD were steeper, and probably closer to the optimal distribution which maximizes photosynthesis, than the average distribution in the stand. Leaves of subordinate plants had a lower mass per unit area (LMA) than those of dominant plants. In the dominant plants, LMA decreased with decreasing relative PFD (and with leaf position) while in the subordinate plants it increased. This surprising result for the subordinate plants can be explained by the fact that, during the course of a growing season, these plants became increasingly shaded and newer leaves were thus formed at progressively lower light availability. This indicates that LMA was strongly determined by the relative PFD at leaf formation and to a lesser extent by the current PFD. Leaf N content per unit mass (Nmass) was strongly determined by leaf position independent of relative PFD. This indicates that Nmass is strongly ontogenetically related to the leaf-aging process while changes in Narea, in response to PFD, were regulated through changes in LMA.

Plant responses to elevated CO2 concentration at different scales: leaf, whole plant, canopy, and population.

Elevated CO2 enhances photosynthesis and growth of plants, but the enhancement is strongly influenced by the availability of nitrogen. In this article, we summarise our studies on plant responses to elevated CO2. The photosynthetic capacity of leaves depends not only on leaf nitrogen content but also on nitrogen partitioning within a leaf. In Polygonum cuspidatum, nitrogen partitioning among the photosynthetic components was not influenced by elevated CO2 but changed between seasons. Since the alteration in nitrogen partitioning resulted in different CO2-dependence of photosynthetic rates, enhancement of photosynthesis by elevated CO2 was greater in autumn than in summer. Leaf mass per unit area (LMA) increases in plants grown at elevated CO2. This increase was considered to have resulted from the accumulation of carbohydrates not used for plant growth. With a sensitive analysis of a growth model, however, we suggested that the increase in LMA is advantageous for growth at elevated CO2 by compensating for the reduction in leaf nitrogen concentration per unit mass. Enhancement of reproductive yield by elevated CO2 is often smaller than that expected from vegetative growth. In Xanthium canadense, elevated CO2 did not increase seed production, though the vegetative growth increased by 53%. As nitrogen concentration of seeds remained constant at different CO2 levels, we suggest that the availability of nitrogen limited seed production at elevated CO2 levels. We found that leaf area development of plant canopy was strongly constrained by the availability of nitrogen rather than by CO2. In a rice field cultivated at free-air CO2 enrichment, the leaf area index (LAI) increased with an increase in nitrogen availability but did not change with CO2 elevation. We determined optimal LAI to maximise canopy photosynthesis and demonstrated that enhancement of canopy photosynthesis by elevated CO2 was larger at high than at low nitrogen availability. We also studied competitive asymmetry among individuals in an even-aged, monospecific stand at elevated CO2. Light acquisition (acquired light per unit aboveground mass) and utilisation (photosynthesis per unit acquired light) were calculated for each individual in the stand. Elevated CO2 enhanced photosynthesis and growth of tall dominants, which reduced the light availability for shorter subordinates and consequently increased size inequality in the stand.

Plants in a crowded stand regulate their height growth so as to maintain similar heights to neighbours even when they have potential advantages in height growth

BACKGROUND AND AIMS Although being tall is advantageous in light competition, plant height growth is often similar among dominant plants in crowded stands (height convergence). Previous theoretical studies have suggested that plants should not overtop neighbours because greater allocation to supporting tissues is necessary in taller plants, which in turn lowers leaf mass fraction and thus carbon gain. However, this model assumes that a competitor has the same potential of height growth as their neighbours, which does not necessarily account for the fact that height convergence occurs even among individuals with various biomass. METHODS Stands of individually potted plants of Chenopodium album were established, where target plants were lifted to overtop neighbours or lowered to be overtopped. Lifted plants were expected to keep overtopping because they intercept more light without increased allocation to stems, or to regulate their height to similar levels of neighbours, saving biomass allocation to the supporting organ. Lowered plants were expected to be suppressed due to the low light availability or to increase height growth so as to have similar height to the neighbours. KEY RESULTS Lifted plants reduced height growth in spite of the fact that they received higher irradiance than others. Lowered plants, on the other hand, increased the rate of stem elongation despite the reduced irradiance. Consequently, lifted and lowered plants converged to the same height. In contrast to the expectation, lifted plants did not increase allocation to leaf mass despite the decreased stem length. Rather, they allocated more biomass to roots, which might contribute to improvement of mechanical stability or water status. It is suggested that decreased leaf mass fraction is not the sole cost of overtopping neighbours. Wind blowing, which may enhance transpiration and drag force, might constrain growth of overtopping plants. CONCLUSIONS The results show that plants in crowded stands regulate their height growth to maintain similar height to neighbours even when they have potential advantages in height growth. This might contribute to avoidance of stresses caused by wind blowing.

Architecture and growth of an annual plant Chenopodium album 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.

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.

Photoperceptive Site in Phytochrome-Mediated Lettuce (Lactuca sativa L. cv. Grand Rapids) Seed Germination

Summary The photoperceptive site in phytochrome-mediated lettuce seed germination was determined using micro-beam irradiation. To avoid the shading effect of black pericarps, decoated seeds were incubated in 0.2 M sorbitol solution. Red light exposed on the second section of the octasection along the longitudinal axis (0.4–0.8 mm) from the root tips could induce as high a germination rate as whole-seed irradiation. Red light on the other parts brought a low germination rate. Anatomical studies suggested that this photosensitive portion corresponded to the hypocotyl, because this part was the upper half of the embryonic axis and it never formed root hairs later. Red light-induced growth at first occurred in the hypocotyl and the root region simultaneously. Therefore, the light signal probably was transferred from hypocotyl to root.

The role of biomass allocation between lamina and petioles in a game of light competition in a dense stand of an annual plant

Background and Aims Models of plant three-dimensional (3-D) architecture have been used to find optimal morphological characteristics for light capture or carbon assimilation of a solitary plant. However, optimality theory is not necessarily useful to predict the advantageous strategy of an individual in dense stands, where light capture of an individual is influenced not only by its architecture but also by the architecture of its neighbours. Here, we analysed optimal and evolutionarily stable biomass allocation between the lamina and petiole (evolutionarily stable strategy; ESS) under various neighbour conditions using a 3-D simulation model based on the game theory. Methods We obtained 3-D information of every leaf of actual Xanthium canadense plants grown in a dense stand using a ruler and a protractor. We calculated light capture and carbon assimilation of an individual plant when it stands alone and when it is surrounded by neighbours in the stand. We considered three trade-offs in petiole length and lamina area: biomass allocation, biomechanical constraints and photosynthesis. Optimal and evolutionarily stable biomass allocation between petiole and lamina were calculated under various neighbour conditions. Key Results Optimal petiole length varied depending on the presence of neighbours and on the architecture of neighbours. The evolutionarily stable petiole length of plants in the stand tended to be longer than the optimal length of solitary plants. The mean of evolutionarily stable petiole length in the stand was similar to the real one. Trade-offs of biomechanical constraint and photosynthesis had minor effects on optimal and evolutionarily stable petiole length. Conclusion Actual plants realize evolutionarily stable architecture in dense stands. Interestingly, there were multiple evolutionarily stable petiole lengths even in one stand, suggesting that plants with different architectures can coexist across plant communities.