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     

Branching, nitrogen distribution, and carbon gain in solitary plants of an annual herb, Xanthium canadense

The theory of optimal nitrogen (N) distribution predicts that the carbon gain of plants will be maximised when leaves of higher irradiance have higher N content per area (N _area). Most previous studies have examined optimal N distribution without explicitly considering the branching status of plants. I investigated light environment, N distribution and photosynthetic traits of individual leaves of an herbaceous species, Xanthium canadense. X. canadense was grown solitary under high (HN) and low nutrients (LN). Light availability, leaf mass per unit area and N _area were measured for all leaves within plants. Daily photosynthesis of the plants of actual N distribution was compared with those of optimal and constant N distribution. Branch production was facilitated in HN but not in LN plants. N _area was correlated more with leaf order than with leaf light environment. Although N was more limited and the light environment was less heterogeneous within crowns in LN than in HN plants, leaf N distribution was closer to optimal in the latter. These results suggest that leaf N distribution was not optimised in solitary plants of X. canadense. Because this species often regenerates in a dense stand, leaf N distribution might be selected to maximise carbon gain only in such a stand. Leaf N distribution might thus be constrained by the regeneration strategy of the species.     

Reproductive yield of individuals competing for light in a dense stand of an annual, Xanthium canadense

In a dense stand, individuals compete with each other for resources, especially for light. Light availability decreases with increasing depth in the canopy, thus light competition becoming stronger with time in the vegetative phase. In the reproductive phase, on the other hand, leaves start senescing, and the light environment, particularly of smaller individuals, will be improved. To study the effect of change in light climate on reproduction of individuals, we established an experimental stand of an annual, Xanthium canadense, and assessed temporal changes in whole plant photosynthesis through the reproductive phase with particular reference to light availability of individuals. At flowering, 83% of individuals were still alive, but only 27% survived to set seeds. Most of the individuals that died in the reproductive phase were smaller than those that produced seeds. Individuals that died at the early stage of the reproductive phase had a lower leaf to stem mass ratio, suggesting that the fate of individuals was determined partly by the pattern of biomass allocation in this period. At the early stage of the reproductive phase, leaf area index (LAI) of the stand was high and larger individuals had higher whole plant photosynthesis than smaller individuals. Although light availability at later stages was improved with reduction in LAI, whole plant photosynthesis was very low in all individuals due to a lower light use efficiency, which was caused by a decrease in photosynthetic N use efficiency. We conclude that light competition was still strong at the early stage of the reproductive phase and that later improvement of light availability did not ameliorate the photosynthesis of smaller individuals.     

Nitrogen use efficiency in instantaneous and daily photosynthesis of leaves in the canopy of a Solidago altissima stand

Photosynthetic capacity was measured on detached leaves sampled in a canopy of Solidago altissima L. Non-rectangular hyperbola fitted the light response curve of photosynthesis and significant correlations were observed between leaf nitrogen per unit area and four parameters which characterize the light-response curve. Using regressions of the parameters on leaf nitrogen, a model of leaf photosynthesis was constructed which gave the relationships between leaf nitrogen, photon flux density (PFD) and photosynthesis. Curvilinear relations were obtained between leaf nitrogen and photosynthetic rate on both an instantaneous and a daily basis. Nitrogen use efficiency (NUE, photosynthesis per unit leaf nitrogen) was calculated against leaf nitrogen under varying PFDs. The optimum nitrogen content per unit leaf area that maximizes NUE shifted to higher values with increasing PFD. Field measurements of PFD showed high positive correlations between the distribution of leaf nitrogen in the canopy and relative PFD. The predicted optimum leaf nitrogen content for each level in the canopy, to achieve maximized NUE during a clear day, was close to the actual nitrogen distribution as found through sampling.     

Estimating photosynthetic radiation use efficiency using incident light and photosynthesis of individual leaves

It has been theorized that photosynthetic radiation use efficiency (PhRUE) over the course of a day is constant for leaves throughout a canopy if leaf nitrogen content and photosynthetic properties are adapted to local light so that canopy photosynthesis over a day is optimized. To test this hypothesis, 'daily' photosynthesis of individual leaves of Solanum melongena plants was calculated from instantaneous rates of photosynthesis integrated over the daylight hours. Instantaneous photosynthesis was estimated from the photosynthetic responses to photosynthetically active radiation (PAR) and from the incident PAR measured on individual leaves during clear and overcast days. Plants were grown with either abundant or scarce N fertilization. Both net and gross daily photosynthesis of leaves were linearly related to daily incident PAR exposure of individual leaves, which implies constant PhRUE over a day throughout the canopy. The slope of these relationships (i.e. PhRUE) increased with N fertilization. When the relationship was calculated for hourly instead of daily periods, the regressions were curvilinear, implying that PhRUE changed with time of the day and incident radiation. Thus, linearity (i.e. constant PhRUE) was achieved only when data were integrated over the entire day. Using average PAR in place of instantaneous incident PAR increased the slope of the relationship between daily photosynthesis and incident PAR of individual leaves, and the regression became curvilinear. The slope of the relationship between daily gross photosynthesis and incident PAR of individual leaves increased for an overcast compared with a clear day, but the slope remained constant for net photosynthesis. This suggests that net PhRUE of all leaves (and thus of the whole canopy) may be constant when integrated over a day, not only when the incident PAR changes with depth in the canopy, but also when it varies on the same leaf owing to changes in daily incident PAR above the canopy. The slope of the relationship between daily net photosynthesis and incident PAR was also estimated from the photosynthetic light response curve of a leaf at the top of the canopy and from the incident PAR above the canopy, in place of that measured on individual leaves. The slope (i.e. net PhRUE) calculated in this simple way did not differ statistically from that calculated using data from individual leaves.     

Relationships between soil microbial properties and aboveground stand characteristics of conifer forests in Oregon

Eight forest sites representing a large range of climate, vegetation, and productivity were sampled in a transect across Oregon to study the relationships between aboveground stand characteristics and soil microbial properties. These sites had a range in leaf area index of 0.6 to 16 m² m^–2 and net primary productivity of 0.3 to 14 Mg ha^–1 yr^–1.Measurements of soil and forest floor inorganic N concentrations and in situ net N mineralization, nitrification, denitrification, and soil respiration were made monthly for one year. Microbial biomass C and anaerobic N mineralization, an index of N availability, were also measured. Annual mean concentrations of NH ₄ ⁺ ranged from 37 to 96 mg N kg^–1 in the forest floor and from 1.7 to 10.7 mg N kg^–1 in the mineral soil. Concentrations of NO ₃ ^– were low ( < 1 mg N kg^–1) at all sites. Net N mineralization and nitrification, as measured by the buried bag technique, were low on most sites and denitrification was not detected at any site. Available N varied from 17 to 101 mg N kg^–1, microbial biomass C ranged from 190 to 1230 mg Ckg^–1, and soil respiration rates varied from 1.3 to 49 mg C kg^–1 day^–1 across these sites. Seasonal peaks in NH ₄ ⁺ concentrations and soil respiration rates were usually observed in the spring and fall.The soils data were positively correlated with several aboveground variables, including leaf area index and net primary productivity, and the near infrared-to-red reflectance ratio obtained from the airborne simulator of the Thematic Mapper satellite. The data suggest that close relationships between aboveground productivity and soil microbial processes exist in forests approaching semi-equilibrium conditions.Abbreviations IR infrared - LAI leaf area index - k _c proportion of microbial biomass C mineralized to CO₂ - NPP net primary productivity - TM Thematic Mapper     

Timing of canopy closure influences carbon translocation and seed production of an understorey herb, Trillium apetalon (Trilliaceae)

BACKGROUND AND AIMS: The light availability on a temperate, deciduous-forest floor varies greatly, reflecting the seasonal leaf dynamics of the canopy trees. The growth and/or reproductive activity of understorey plants should be influenced by the length of the high-irradiance period from snowmelt to canopy closure. The aim of the present study was to clarify how spring-blooming species regulate the translocation of photosynthetic products to current reproduction and storage organs during a growing season in accordance with the changing light conditions. METHODS: Growth pattern, net photosynthetic rate, seed production, and shoot and flower production in the next year of Trillium apetalon were compared between natural and experimentally shaded conditions. Furthermore, translocation of current photosynthetic products within plants was assessed by a labelled carbon-chase experiment. KEY RESULTS: During the high-irradiance period, plants showed high photosynthetic ability, in which current products were initially used for shoot growth, then reserved in the rhizome. Carbon translocation to developing fruit occurred after canopy closure, but this was very small due to low photosynthetic rates under the darker conditions. The shading treatment in the early season advanced the time of carbon translocation to fruit, but reduced seed production in the current year and flower production of the next year. CONCLUSIONS: Carbon translocation to the storage organ had priority over seed production under high-irradiance conditions. A shortened bright period due to early canopy closure effectively restricts carbon assimilation, which greatly reduces subsequent reproductive output owing to low photosynthetic products for fruit development and small carbon storage for future reproduction. As populations of this species are maintained by seedling recruitment, acceleration of canopy closure timing may influence the maintenance and dynamics of populations.     

Leaf area index, canopy structure and photosynthesis of red clover (Trifolium pratense L.)

Abstract. The influence of leaf age, total leaf area and its dispersion in space on canopy photosynthesis were studied using microswards of red clover ( Trifolium pratense L.) which were established in the greenhouse. Two varieties, Renova (flowering) and Molstad (non-flowering), were sown in separate plastic boxes at densities of 225, 400 and 625 plants per m².
Vertical distribution of photosynthetically active radiation (PAR), leaf area, leaf age and ¹⁴CO₂-fixation were determined periodically. Net photosynthesis and dark respiration of canopies were measured. Maximum photosynthetic capacity of individual leaves was measured on plants taken from the intact canopy or from plants where shading of the growing leaves had been prevented.
Net photosynthetic rate of canopies increased linearly with leaf area index (LAI) up to an LAI of 3.5 and then declined at higher LAI, independent of variety and sowing density. Below the optimum LAI, net photosynthesis depended mainly on interception of PAR. Decrease in canopy photosynthesis above the optimum LAI was due to a higher proportion of old leaves with decreased photosynthetic capacity, and not to an increase in respiring plant parts. It is concluded that LAI and position of leaf age categories in the canopy are more important than vertical distribution of leaf area in determining canopy photosynthesis of red clover.     

Scaling sun and shade photosynthetic acclimation of Alocasia macrorrhiza to whole-plant performance – II. Simulation of carbon balance and growth at different photon flux densities

A whole-plant carbon balance model incorporating a light acclimation response was developed for Alocasia macrorrhiza based on empirical data and the current understanding of light acclimation in this species. The model was used to predict the relative growth rate (RGR) for plants that acclimated to photon flux density (PFD) by changing their leaf type, and for plants that produced only sun or shade leaves regardless of PFD. The predicted RGR was substantially higher for plants with shade leaves than for those with sun leaves at low PFD. However, the predicted RGR was not higher, and in fact was slightly lower, for plants with sun leaves than for those with shade leaves at high PFD. The decreased leaf area ratios (LARs) of the plants with sun leaves counteracted their higher photosynthetic capacities per unit leaf area (A_max). The model was manipulated by changing parameters to examine the sensitivity of RGR to variation in single factors. Overall, RGR was most sensitive to LAR and showed relatively little sensitivity to variation in A_max or maintenance respiration. Similarly, RGR was relatively insensitive to increases in leaf life-span beyond those observed. Respiration affected RGR only at low PFD, whereas A_max was moderately important only at high PFD.     

Growth and carbon economy of a fast-growing and a slow-growing grass species as dependent on nitrate supply

In previous experiments systematic differences have been found in the morphology, carbon economy and chemical composition of seedlings of inherently fast- and slow-growing plant species, grown at a non-limiting nutrient supply. In the present experiment it was investigated whether these differences persist when plants are grown at suboptimal nutrient supply rates. To this end, plants of the inherently fast-growing Holcus lanatus L. and the inherently slow-growing Deschampsia flexuosa (L.) Trin. were grown in sand at two levels of nitrate supply. Growth, photosynthesis, respiration and carbon and nitrogen content were studied over a period of 4 to 7 weeks. At low N-supply, the potentially fast-growing species still grew faster than the potentially slow-growing one. Similarly, differences in leaf area ratio (leaf area:total dry weight), specific leaf area (leaf area:leaf dry weight) and leaf weight ratio (leaf dry weight:total dry weight), as observed at high N-supply persisted at low N-availability. The only growth parameter for which a substantial Species × N-supply interaction was found was the net assimilation rate (increase in dry weight per unit leaf area and time). Rates of photosynthesis, shoot respiration and root respiration, expressed per unit leaf, shoot and root weight, respectively, were lower for the plants at low N-availability and higher for the fast-growing species. Species-specific variation in the daily carbon budget was mainly due to variation in carbon fixation. Lower values at low N were largely determined by both a lower C-gain of the leaves and a higher proportion of the daily gain spent in root respiration. Interspecific variation in C-content and dry weight:fresh weight ratio were similar at low and high N-supply. Total plant organic N decreased with decreasing N-supply, without differences between species. It is concluded that most of the parameters related to growth, C-economy and chemical composition differ between species and/or are affected by N-supply, but that differences between the two species at high N-availability persist at low N-supply.     

Quantitative analysis of carbon balance in the reproductive organs and leaves of <Emphasis Type="Italic">Cinnamomum camphora</Emphasis> (L.) Presl

We investigated seasonal changes in dry mass and CO₂ exchange rate in fruit and leaves of the evergreen tree Cinnamomum camphora with the aim of quantitatively determining the translocation balance between the two organs. The fruit dry mass growth peaked in both August and October: the first increase was due to fruit pulp development and the second to seed development. Fruit respiration also increased with the rapid increase in fruit dry mass. Therefore, the carbohydrates required for fruit development showed two peaks during the reproductive period. Fruit photosynthesis was relatively high in early August, when fruit potentially re-fixed 75% of respired CO₂, indicating that fruit photosynthesis contributed 15–35% of the carbon requirement for fruit respiration. Current-year leaves completed their growth in June when fruit growth began. Current-year leaves translocated carbohydrates at a rate of approximately 10–25 mg dry weight (dw) leaf⁻¹ day⁻¹ into other organs throughout the entire fruit growth period. This rate of translocation from current-year leaves was much higher than the amount of carbohydrate required for reproduction (ca. 3 mg dw fruit⁻¹ day⁻¹). Given the carbon balance between fruit and current-year leaves, carbohydrates for reproduction were produced within the current-year fruit-bearing shoots. C. camphora would be adaptive for steadily supplying enough amount of carbohydrate to the fruits, as there was little competition for carbohydrates between the two organs. As assimilates by leaves are used for processes such as reproduction and the formation of new shoots, photosynthesis by reproductive organs is considered to be important to compensate for reproductive cost.     

Photosynthesis, respiration, and chlorophylls in presenescent, regreened, and senescent leaves of forest herb Smyrnium perfoliatum L. (Apiaceae)

Smyrnium perfoliatum L. (Apiaceae), an endangered forest herb with only one main locality in South-West Slovakia (Devínska Kobyla in the Little Carpathian Mountains), is capable to recover chlorophylls at the end of the growing season. This regreening only within bracts was observed during two weeks before achieving the so-called “point of no return” that leads to the last stage of ontogenesis — leaf senescence. The effect not only of endogenous cytokinins on chlorophyll content and carbon dioxide exchange (photosynthesis, mitochondrial respiration, and photorespiration) is discussed but also of other factors such as strong irradiance, high temperature or drought stress on studied parameters was considered.     

基于DSSAT模型的吉林省黑土作物-土壤氮循环和土壤有机碳平衡

应用DSSAT模型中的CERES-Maize作物模型和Century土壤模型,分析了作物管理参数、施肥量、土壤初始氮含量和作物桔杆还田对吉林省黑土地区玉米生长、氮循环以及有机碳氮生态平衡的影响.结果表明:在玉米目标产量为12000～15000kg.hm-2条件下,最佳施氮肥量为200～240kgN.hm-2.在该氮肥用量下,玉米地上氮吸收量为250～290kgN.hm-2,其中,120～140kgN.hm-2来自土壤,130～150kgN.hm-2来自肥料;提高氮肥用量(250～420kgN.hm-2)将导致土壤残留氮明显增加(63～183kgN.hm-2);延迟追肥时间同样导致土壤残留氮增加;当玉米秸杆还田量超过6000kg.hm-2时,模拟的土壤活性有机碳、氮可以维持当年的供需平衡.建议在吉林省中部地区黑土玉米带,化肥施氮量控制在200～240kgN.hm-2,适时追肥,秸杆还田量在6000kg.hm-2以上,以确保高产和维持土壤养分生态平衡.     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

Optimal photosynthetic characteristics of individual plants in vegetation stands and implications for species coexistence

AIMS: This paper reviews the way optimization theory has been used in canopy models to analyse the adaptive significance of photosynthesis-related plant characteristics and their consequences for the structure and species composition of vegetation stands. SCOPE: In most studies simple optimization has been used with trait values optimal when they lead to maximum whole-stand photosynthesis. This approach is subject to the condition that the optimum for one individual is independent of the characteristics of its neighbours. This seems unlikely in vegetation stands where neighbour plants strongly influence each other's light climate. Not surprisingly, there are consistent deviations between predicted plant traits and real values: plants tend to be taller, distribute nitrogen more evenly among their leaves and produce more leaf area which is projected more horizontally than predicted by models. CONCLUSIONS: By applying game theory to individual plant-based canopy models, other studies have shown that optimal vegetation stands with maximum whole-stand photosynthesis are not evolutionarily stable. They can be successfully invaded by mutants that are taller, project their leaves more horizontally or that produce greater than optimal leaf areas. While these individual-based models can successfully predict the canopy structure of vegetation stands, they are invariably determined at unique optimal trait values. They do not allow for the co-existence of more than one species with different characteristics. Canopy models can contribute to our understanding of species coexistence through (a) simultaneous analysis of the various traits that determine light capture and photosynthesis and the trade-offs between them, and (b) consideration of trade-offs associated with specialization to different positions in the niche space defined by temporal and spatial heterogeneity of resources.     

植物叶片解剖结构的量化及其在C4植物高粱中的应用

测量切片内所有细胞或细胞器的总周长,并乘以切片厚度和校正因子是一种量化植物叶片解剖结构的常用方法。前人在使用该方法时因忽视了细胞大小、切片厚度等影响估算精确度的因素,从而导致计数样本偏少并产生较大误差。通过分析该方法误差产生的原因及其与精确度的关系,明确了该方法在实际应用中不同精确度所要求对应的切片数量。在此基础上,将该方法用于C4植物高粱(Sorghum bicolor)叶片结构的量化,提出叶肉细胞与维管束鞘细胞接触面积(CABM)等参数,发展并完善了植物叶片解剖结构的量化方法。