Light acquisition and use by individuals competing in a dense stand of an annual herb, Xanthium canadense

The importance of light acquisition and utilization by individuals in intraspecific competition was evaluated by determining growth and photosynthesis of individual plants in a dense monospecific stand of an annual, Xanthium canadense. Photosynthesis of individual plants in the stand was calculated using a canopy photosynthesis model in which leaf photosynthesis was assumed to be function of leaf nitrogen content and light availability. The estimated photosynthetic rates of individuals were strongly correlated with the measured growth rates. Photosynthetic rates per unit aboveground mass (RPR, relative photosynthetic rate) increased with increasing aboveground mass, suggesting asymmetric (one-sided) competition in the stand. However, larger individuals had similar RPRs, suggesting symmetric (two-sided) competition. These results were consistent with the observation that size inequality over the whole stand increased with growth, but it remained stable among the larger individuals. The RPR of an individual was calculated as the product of absorbed photon flux per unit aboveground mass (Φ_mass) and light use efficiency (LUE, photosynthesis per unit absorbed photon flux). Φ_mass indicates the efficiency of light acquisition, and was higher in larger individuals in the stand, while LUE was highest in individuals with intermediate aboveground mass. LUE depends on leaf nitrogen content. At an early stage, leaf nitrogen contents of smaller individuals were similar to those that maximize LUE. Light availability to smaller individuals decreased as they grew, while their nitrogen contents did not change markedly, which decreased their LUE. We concluded that asymmetric competition among individuals in the stand resulted mainly from lower efficiencies in both light acquisition and light use by smaller individuals. Received: 31 January 1998 / Accepted: 12 November 1998     

Plant responses to elevated CO<Subscript>2</Subscript> 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.     

Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N_area) were determined for individuals, and leaf photosynthesis was measured as a function of N_area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P_mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P_mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P_mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P_mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N_area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N_area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.     

Structural and physiological sexual dimorphism estimated from three‐dimensional virtual trees of yerba‐mate (Ilex paraguariensis) is modified by cultivation environment

Yerba‐mate is a subtropical, evergreen, dioecious, South American tree. Sexual dimorphism in photosynthesis, leaf allometry and foliage distribution was hypothesised. Virtual trees (constructed in VPlants software from detailed measurements of plant morphogenesis) of the two genders were compared considering two contrasted cultivation environments and three developmental stages. The total crown volume, leaf area per plant (LA), leaf area index (LAI) and leaf area density (LAD) were calculated. The light interception and photosynthesis were computed from mock‐ups in VegeSTAR. Structural sexual dimorphism concerned general plant form, internode length, leaf allometry, leaf surface, pattern of leaf area distribution and LAD. Cultivation environment and developmental stage acted strongly on sex expression of all observed structural parameters and physiological stages. Sexual differentiation in LA and light interception was related to leaves positioned in the lowest layers (150 cm above ground), whereas sexual specialisation in leaf and plant photosynthesis was related to early vegetative and reproductive stages. Several sexual responses strongly depended on the environment, especially light conditions, with opposite effects observed on female and male plants whether they were cultivated in monoculture or in forest understorey, under high‐light condition or low‐light condition, respectively. Optimised foliage structure and physiology in females may compensate for greater reproductive costs in early developmental stages, but females and males equalise in photosynthetic efficiency after 2‐year regrowth.     

Evolutionarily stable leaf area production in plant populations

Using an analytical model, it was shown that for a given amount of nitrogen in the canopy of a stand (N(T)), there exists an evolutionarily stable leaf area index (ES-LAI), and therefore an evolutionarily stable average leaf nitrogen content (n(ES)(av);n(ES)(av) =N(T)/ES-LAI), at which no individual plant in the stand can increase its photosynthesis by changing its leaf area. It was also shown that this ES-LAI is always greater than the optimal LAI that maximizes photosynthesis per unit N(T) of the stand. This illustrates that the canopy structure that maximizes photosynthesis of a population is not the same as the canopy structure that maximizes photosynthesis of individuals within a population. It was further derived that the ES-LAI at given N(T) increases with the ratio between the light-saturated photosynthesis and the N content per unit leaf area (leaf-PPNUE) and that it decreases with the canopy extinction coefficient for light (K(L)), the light availability and the apparent quantum yield (phi). These hypotheses were tested by comparing calculated ES-LAI and n(ES)(av) values to actual LAIs and leaf N contents measured for stands of a large variety of herbaceous plants. There was a close correspondence between the calculated and measured values. As predicted by the model, plants with high leaf-PPNUEs produced more leaf area per unit nitrogen than those with low leaf-PPNUEs while plants with horizontal leaves, forming stands with higher K(L) values, produced less leaf area than those with more vertically inclined leaves. These results suggest that maximization of individual plant photosynthesis per unit of nitrogen plays an important role in determining leaf area production of plants and the resulting canopy structure of stands of vegetation. They further suggest this optimization to be a mechanism by which leaf traits such as leaf-PPNUE and leaf inclination angle are causally related to structural characteristics of the population, i.e. the leaf area index of the stand.     

Development of the Monsi-Saeki theory on canopy structure and function

BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.     

Life history and population dynamics of the Japanese Yam,Dioscorea japonica thunb.

Shoot morphology at the emergence ofDioscorea japonica Thumb. could be classed into the following three types: (1) a seedling emergence with only one leaf (Se type), (2) a plant consisting of one stem and one leaf, which has emerged from a small tuber (rhizophore) or bulbil less than 50 mg in dry weight (S type) and (3) a twiner with many leaves, which has emerged from a tuber or a bulbil of more than 50 mg in dry weight (L type). The Se type failed to develop beyond the second leaf stage in 1.5% sunlight exposure. The effects of initial plant (seeds, bulbils and tubers) size and light intensity on plant growth were analyzed. The larger the initial plant size was, the greater the growth in height and leaf area was. The distribution ratio of assimilated substances in leaves was high in smaller plants at the early growth stage. The distribution ratio in the tubers of larger plants became high at the early stage of growth. In all three types at over 3% sunlight exposure, the switch-over from the vegetative to reproductive growth phase occurred simultaneously at a later growth stage, but the Se type at 1.5% sunlight exposure showed a very early switch-over in its development; this switch-over may be related to shade tolerance capacity. The L type showed shade avoidance by forming a large productive structure as a twiner     

Canopy structure,nitrogen distribution and whole canopy photosynthetic carbon gain in growing and flowering stands of tall herbs

Using a combination of mathematical modeling and field studies we showed that in dense stands of growing herbaceous plants the vertical pattern of leaf nitrogen distribution resembles the pattern of mean light attenuation in the stand and hence tends to maximize total daily photosynthetic carbon gain of the whole stand. Flowering represents a strong sink of nitrogen away from the photosynthetic apparatus and in herbs like Solidago altissima it induces leaf shedding. We studied both the effect of nitrogen reallocation and leaf shedding on the whole canopy photosynthesis and changes in leaf nitrogen distributions in stands moving from the growing to the flowering stage. Despite a decrease in leaf area index and total nitrogen available for photosynthesis in the flowering stand, the leaf nitrogen distribution here also leads to an almost maximum canopy photosynthesis. In both the growing and the flowering stands the leaf area index was higher than calculated optimum values. It is pointed out that this should not necessarily be interpreted as non-adaptive.     

Growth of Individuals in Plant Populations

Relationships between individual plant weight and net photosynthesisper day (G(t, x) function of plant weight) in plant populationsof various stand structures were simulated based on a canopyphotosynthesis model. The G(t, x) functions of plant weightare determined mainly by LAI (leaf area index), the relationshipbetween individual plant weight and leaf area, canopy structureand extinction coefficient. The concave relationship betweenindividual plant weight and leaf area at small LAI (<2),at small extinction coefficient (< 0.5), or at the canopystructure having the maximum leaf area density at the bottomproduces a concave G(t, x) function, which generates negativeskewness of plant weight. The linear relationship between individualplant weight and leaf area at large LAI (> 2) produces aconvex G(t, x) function, which generates positive skewness ofplant weight. These simulation results coincide with G(t, x)functions obtained experimentally and with the well-known phenomenonof stand dynamics in which skewness of plant weight becomesnegative in the early growth stage and then increases to a positivevalue as a stand grows and becomes crowded. Helianthus annuus L., individual plant size, mean growth rate, canopy photosynthesis model, skewness, stand structure     

Population distributions of plant size and light environment of giant ragweed (Ambrosia trifida L.) at three densities

Summary Plots in a naturally occurring population of giant ragweed (Ambrosia trifida L.) near Ames, Iowa, USA were left unthinned (high density,=693 plants/m²) or were thinned in early June 1989 to create low and medium densities of 10 and 50 plants/m². Size and light environment of individual plants were measured at monthly intervals from June to September. By September, low density plants had 15 times greater biomass/plant and 30 times greater leaf area/plant than high density plants, although biomass and leaf area per unit land area decreased with decreasing density. Plants at high density allocated more biomass to stem growth, but plants at medium and low density had successively higher leaf area ratios, higher potential photosynthetic rates, higher allocation to leaves, and higher growth rates. Average light on leaves decreased with increasing density and also decreased over the growing season in the low and medium densities. The distribution of light environments of individual plants was non-normal and skewed to the left in most months, in contrast to the rightwards skew of distributions of plant size parameters. Inequality in the distributions, as measured by coefficient of variation and Gini coefficients, increased over most of the growing season. There was little effect of density on inequality of stem diameter, height, or estimated dry weight, but inequality in reproductive output greatly increased with density. There was greater inequality in number of staminate flowers produced than in number of pistillate flowers and seeds produced. Path analysis indicated that early plant size was the most important predictor of final plant size and reproductive output; photosynthesis, conductance, and light environment were also significantly correlated with size and reproduction but usually were of minor importance. Variation in growth rate apparently increased inequality in plant size at low density, whereas belowground competition and death of smaller plants may have limited increases in inequality at high density.     

Seasonal changes in photosynthesis of four understory herbs in deciduous forests

Seasonal patterns of photosynthesis and respiration of single leaves of four understory perennial herbs in deciduous forests were investigated in relation to their leaf growth and light conditions on the forest floor.Anemone flaccida shows rapid growth of leaf area and high rates of gross photosynthesis at light saturation (Psat) in its early stage of development. Its photosynthetic activity is restricted to a brief period of high light intensity before the closure of overstory canopies.Disporum smilacinum possesses light-photosynthesis curves of the shade-leaf type throughout its whole growing period. A shading experiment has shown that this plant is low-light adapted and can utilize weak light efficiently. The light-photosynthesis curve ofSyneilesis palmata shifts from the sun-leaf type to the shade-leaf type in response to the seasonal change of light regime on the forest floor. Evergreen leaves ofPyrola japonica have three year longevity, and light-photosynthesis curves of the shade-leaf type. They maintain some photosynthetic activity even in late autumn and winter.     

Plasticity of reproductive traits responding to variation in light availability at the rosette stage of the first year in a strict biennial,Pedicularis torta,from a field on the Qinghai–Tibet Plateau,China

Light is a fundamental heterogeneous environmental factor for plants and often varies at both spatial and temporal scales. In the present study, we carried out a shading experiment to investigate the effect of variable light availability at the rosette stage of the first year in the strict biennial species Pedicularis torta (an endemic species of the Qinghai‐Tibet Plateau) on reproductive traits (i.e. total number of flowers per plant, total number of fruits per plant/total number of flowers per plant, the number of seeds per fruit, mean individual seed mass and estimation of the total number of seeds per plant, plant size at maturity in the second year and on the relationships between reproductive traits and plant size at this stage). We found that light availability affected most of the reproductive traits (except for mean individual seed mass), plant size at maturity in the second year, and the relationship between the total number of flowers per plant and plant size at this stage. Our results indicate that reproductive traits respond to changes in light availability during the rosette plant stage.     

藜个体在高密度种群中的氮素利用效率

 氮素利用效率（NUE）是植物养分策略研究中的一项重要内容。该文利用Berendse和Aerts提出的氮素利用效率概念和原理研究了高密度的藜(Chenopodium album)种群中不同植物个体在种内竞争条件下的氮素利用效率。结果表明,由于植株的氮素吸收速率与其个体大小成非线性关系,说明不同植株个体对氮素的竞争属于非对称竞争。个体较大的植株氮素输入较高,而个体较小的植株氮素输出较高,因而较大个体植株的氮素净增加也较高。植株的氮素损失随着个体大小的增加而增加,较大植株个体的氮素浓度随着生长而下降,而较小植株个体的氮素浓度随时间的变化不大,说明个体较小的植株的生长受光照的限制比受氮素的限制更大,而对较大的植株个体而言,它们的生长受氮素的限制更大。高密度藜种群中的不同植物个体具有不同的养分策略,氮素利用效率及其组成部分氮素生产力（NP）和氮素滞留时间（MRT）均不同。植株的NP和MRT与其个体大小正相关,较大的植物个体具有较高的NP和较长的MRT,因而氮素利用效率也高于个体较小的植株。在个体水平上,种内不同植株的NP与MRT不存在权衡关系（Trade-off）。因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分策略,对于研究种内不同植株的养分策略也同样适用。     

Leaf lifespan and lifetime carbon balance of individual leaves in a stand of an annual herb, Xanthium canadense

Leaf lifespan in response to resource availability has been documented in many studies, but it still remains uncertain what determines the timing of leaf shedding. Here, we evaluate the lifetime carbon (C) balance of a leaf in a canopy as influenced by nitrogen (N) availability. Stands of Xanthium canadense were established with high-nitrogen (HN) and low-nitrogen (LN) treatments and temporal changes of C gain of individual leaves were investigated with a canopy photosynthesis model. Daily C gain of a leaf was maximal early in its development and subsequently declined. Daily C gain at shedding was nearly zero in HN, while it was still positive in LN. Sensitivity analyses showed that the decline in the daily C gain resulted primarily from the reduction in light level in HN and by the reduction in leaf N in LN. Smaller leaf size in LN than in HN led to higher light levels in the canopy, which helped leaves of the LN stand maintain for a longer period. These results suggest that the mechanism by which leaf lifespan is determined changes depending on the availability of the resource that is most limiting to plant growth.     

马铃薯/大豆套作对3个大豆品种光合指标和产量的影响

以大豆品种中黄30(早熟)、冀豆17(中熟)和齐黄34(晚熟)单作为对照,在大田条件下,研究马铃薯/大豆套作模式中3个品种生育期叶面积指数的变化及干物质积累的特征,分析套作马铃薯收获前后对大豆光合指标、产量及其构成因素的影响.结果表明： 生育前期阴蔽导致套作大豆叶面积指数(LAI)降低,干物质积累变缓,营养生长期相对延长,不同品种套作大豆光合速率(P_n)、蒸腾速率(T_r)和气孔导度(g_s)均低于单作.生育后期套作大豆生长加快,尤其是马铃薯收获后晚熟品种增幅显著提高.此时,套作大豆受光条件得到较大改善,表现出较强的补偿效应,LAI、干物质积累、P_n和g_s相对于单作上升幅度加大,接近于单作大豆,但不同品种补偿能力不同.与单作相比,套作模式下不同大豆品种的有效荚数、单株粒数及每荚粒数均有所降低,其中早熟品种分别显著下降22.0%、36.0%、17.6%,中熟品种下降5.1%、13.1%、8.9%,晚熟品种下降5.7%、7.6%、2.1%.套作模式下,中、晚熟大豆品种的产量较早熟品种分别高92.4%和163.4%,总土地当量比(LER)分别达到1.81和1.84.表明中、晚熟大豆品种与马铃薯组合套作优势更强,有利于马铃薯收获后大豆的补偿生长,促进套作大豆产量提高,充分发挥了复合群体的产量优势.     

Effect of Canopy Structure on Degree of Asymmetry of Competition in Two Forest Stands in Northern Japan

The canopy structure in terms of the vertical distribution of leaf mass and the degree of asymmetry of competition between individual trees was studied in two types of forest stand in Hokkaido, northern Japan: a naturally regenerated stand ofBetulaspp. and an artificial plantation ofPicea abies.The canopy structure in theBetulastand was more hierarchical; larger individuals were not heavily shaded even in the lowest part of their crowns and smaller individuals were heavily shaded by their larger neighbours. The canopy structure in thePiceastand was less hierarchical; even larger individuals were shaded in the lowest part of their crowns and smaller individuals were not heavily shaded by their neighbours. Application of the general formula of size-dependent mean growth rate revealed that competition in theBetulastand was more one-sided than that in thePiceastand. This result was consistent with the trends in the change over time in size equality in both stands.Even if competition is mediated by light, which often makes competition one-sided, the degree of one-sidedness in competition can be variable depending on canopy structure.     

Physiological N response of field-grown maize hybrids (Zea mays L.) with divergent yield potential and grain protein concentration

Fertilizer N availability impacts photosynthesis and crop performance, although cause–effect relationships are not well established, especially for field-grown plants. Our objective was to determine the relationship between N supply and photosynthetic capacity estimated by leaf area index (LAI) and single leaf photosynthesis using genetically diverse field-grown maize (Zea mays L.) hybrids. We compared a high yield potential commercial hybrid (FR1064 x LH185) and an experimental hybrid (FR1064 x IHP) with low yield potential but exceptionally high grain protein concentration. Plant biomass and physiological traits were measured at tassel emergence (VT) and at the grain milk stage (R3) to assess the effects of N supply on photosynthetic source capacity and N uptake, and grain yield and grain N were measured at maturity. Grain yield of FR1064 x LH185 was much greater than FR1064 x IHP even though plant biomass and LAI were larger for FR1064 x IHP, and single leaf photosynthesis was similar for both hybrids. Although photosynthetic capacity was not related to hybrid differences in productivity, increasing N supply led to proportional increases in grain yield, plant biomass, LAI, photosynthesis, and Rubisco and PEP carboxylase activities for both hybrids. Thus, a positive relationship between photosynthetic capacity and yield was revealed by hybrid response to N supply, and the relationship was similar for hybrids with a marked difference in yield potential. For both hybrids the N response of single leaf CER and initial Rubisco activity was negative when expressed per unit of leaf N. In contrast, PEP carboxylase activity per unit leaf N increased in response to N availability, indicating that PEP carboxylase served as a reservoir for excess N accumulation in field-grown maize leaves. The correlation between CER and initial Rubisco activity was highly significant when expressed on a leaf area or a total leaf basis. The results suggest that regardless of genotypic yield potential, maize CER, and potentially grain yield, could be improved by increasing the partitioning of N into Rubisco.     

Carbon balance in a monospecific stand of an annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis> at an elevated CO<Subscript>2</Subscript> concentration

Elevated CO₂ enhances carbon uptake of a plant stand, but the magnitude of the increase varies among growth stages. We studied the relative contribution of structural and physiological factors to the CO₂ effect on the carbon balance during stand development. Stands of an annual herb Chenopodium album were established in open-top chambers at ambient and elevated CO₂ concentrations (370 and 700 μmol mol⁻¹). Plant biomass growth, canopy structural traits (leaf area, leaf nitrogen distribution, and light gradient in the canopy), and physiological characteristics (leaf photosynthesis and respiration of organs) were studied through the growing season. CO₂ exchange of the stand was estimated with a canopy photosynthesis model. Rates of light-saturated photosynthesis and dark respiration of leaves as related with nitrogen content per unit leaf area and time-dependent reduction in specific respiration rates of stems and roots were incorporated into the model. Daily canopy carbon balance, calculated as an integration of leaf photosynthesis minus stem and root respiration, well explained biomass growth determined by harvests (r ² = 0.98). The increase of canopy photosynthesis with elevated CO₂ was 80% at an early stage and decreased to 55% at flowering. Sensitivity analyses suggested that an alteration in leaf photosynthetic traits enhanced canopy photosynthesis by 40–60% throughout the experiment period, whereas altered canopy structure contributed to the increase at the early stage only. Thus, both physiological and structural factors are involved in the increase of carbon balance and growth rate of C. album stands at elevated CO₂. However, their contributions were not constant, but changed with stand development.     

Effects of Physiological and Environmental Variations on Size-Structure Dynamics in Plant Populations

Sensitivity analysis was conducted, based on the canopy photosynthesisand continuity equation models which were developed in a previouspaper (Yokozawa and Hara, 1992), to investigate effects of variationin physiological parameters (maximal photosynthetic rate perunit leaf area, respiration rate per unit leaf area, maintenancerespiration rate per unit weight, growth respiration rate perunit weight, light extinction coefficient of the canopy, etc.)on the size-structure dynamics in plant populations. As thedegree of asymmetry in competition between individuals increased,effects of variation in physiological parameters diminished.Therefore, a population undergoing one-sided competition (mostasymmetric competition) is a stable system, little affectedby temporal and spatial variations in the environmental conditionswhich lead to variation in physiological parameters, whereasa population undergoing symmetric two-sided competition is sensitiveto these fluctuations. It was also shown by simulation thatthe degree of asymmetry in competition decreases (through effectson canopy photosynthesis) as nutrient level in the soil is reduced.It is suggested that symmetric two-sided competition is associatedwith non-transitivity of competition between species (i.e. frequentreversals of rank order of species), and hence with speciesdiversity. Several other ecological phenomena are discussedin relation to allometry (i.e. allocation-growth pattern) andthe degree of asymmetry in competition.Copyright 1994, 1999Academic Press Allometry, canopy photosynthesis, competition mode, continuity equation, parameter sensitivity, stability of stand structure     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.