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.     

Canopy structure and leaf nitrogen distribution in a stand of Lysimachia vulgaris L. as influenced by stand density

Summary A hypothesis that a dense stand should develop a less uniform distribution of leaf nitrogen through the canopy than an open stand to increase total canopy photosynthesis was tested with experimentally established stands of Lysimachia vulgaris L. The effect of stand density on spatial variation of photon flux density, leaf nitrogen and specific leaf weight within the canopy was examined. Stand density had little effect on the value of the light extinction coefficient, but strongly affected the distribution of leaf nitrogen per unit area within a canopy. The open stand had more uniform distribution of leaf nitrogen than the dense stand. However, different light climates between stands explained only part of the variation of leaf nitrogen in the canopy. The specific leaf weight in the canopy increased with increasing relative photon flux density and with decreasing nitrogen concentration.     

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     

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.     

Optimal leaf area indices in C3 and C4 mono- and dicotyledonous species at low and high nitrogen availability

From an analytical model it was shown that for a given total amount of nitrogen in the canopy, there exists an optimal leaf area index (LAI), and therefore an optimal average leaf introgen content, at which canopy photosynthesis is maximal. If the LAI is increased above this optimum, increased light interception will not compensate for reduction in photosynthetic capacity of the canopy resulting from reduced leaf nitrogen contents. It was further derived from the model that the value of the optimal LAI increases with the photosynthetic nitrogen use efficiency (PNUE) and decreases with the canopy extinction coefficient for light (K_L) and incident photon flux density (PFD) at the top of the canopy. These hypotheses were tested on dense stands of species with different photosynthetic modes and different architectures. A garden experiment was carried out with the C₄ monocot sorghum ( Sorghum bicolor [L.] Moensch cv. Pioneer), the C₃ monocot rice ( Oryza sativa L. cv. Araure 4), the C₄ dicot amaranth ( Amaranthus cruentus L. cv. K113) and the C₃ dicot soybean ( Glycine max [L.] Merr. cv. Williams) at two levels of nitrogen availability.
The C₄ species had higher PNUEs than the C₃ species while the dicots formed stands with higher extinction coefficients for light and had lower PNUEs than the monocots. The C₄ and monocot species were found to have formed more leaf area per unit leaf nitrogen (i.e., had lower leaf nitrogen contents) than the C₃ and dicot species, respectively. These results indicate that the PNUE and the extinction coefficient for light are important factors determining the amount of leaf area produced per unit nitrogen as was predicted by the model.     

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.     

Maximizing daily canopy photosynthesis with respect to the leaf nitrogen allocation pattern in the canopy

Summary A model of daily canopy photosynthesis was constructed taking light and leaf nitrogen distribution in the canopy into consideration. It was applied to a canopy of Solidago altissima. Both irradiance and nitrogen concentration per unit leaf area decreased exponentially with increasing cumulative leaf area from the top of the canopy. The photosynthetic capacity of a single leaf was evaluated in relation to irradiance and nitrogen concentration. By integration, daily canopy photosynthesis was calculated for various canopy architectures and nitrogen allocation patterns. The optimal pattern of nitrogen distribution that maximizes the canopy photosynthesis was determined. Actual distribution of leaf nitrogen in the canopy was more uniform than the optimal one, but it realized over 20% more photosynthesis than that under uniform distribution and 4.7% less photosynthesis than that under the optimal distribution. Redeployment of leaf nitrogen to the top of the canopy with ageing should be more effective in increasing total canopy photosynthesis in a stand with a dense canopy than in a stand with an open canopy.     

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     

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.     

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.     

Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest

Canopy structure and light interception were measured in an 18-m tall, closed canopy deciduous forest of sugar maple (Acer saccharum) in southwestern Wisconsin, USA, and related to leaf structural characteristics, N content, and leaf photosynthetic capacity. Light attenuation in the forest occurred primarily in the upper and middle portions of the canopy. Forest stand leaf area index (LAI) and its distribution with respect to canopy height were estimated from canopy transmittance values independently verified with a combined leaf litterfall and point-intersect method. Leaf mass, N and A _max per unit area (LMA, N/area and A _max/area, respectively) all decreased continuously by over two-fold from the upper to lower canopy, and these traits were strongly correlated with cumulative leaf area above the leaf position in the canopy. In contrast, neither N concentration nor A _max per unit mass varied significantly in relation to the vertical canopy gradient. Since leaf N concentration showed no consistent pattern with respect to canopy position, the observed vertical pattern in N/area is a direct consequence of vertical variation of LMA. N/area and LMA were strongly correlated with A _max/area among different canopy positions (r²=0.81 and r²=0.66, respectively), indicating that vertical variation in area-based photosynthetic capacity can also be attributed to variation in LMA. A model of whole-canopy photosynthesis was used to show that observed or hypothetical canopy mass distributions toward higher LMA (and hence higher N/area) in the upper portions of the canopy tended to increase integrated daily canopy photosynthesis over other LMA distribution patterns. Empirical relationships between leaf and canopy-level characteristics may help resolve problems associated with scaling gas exchange measurements made at the leaf level to the individual tree crown and forest canopy-level.     

美国黑核桃实生苗生态生理过程对环境因素响应的数值模拟(Ⅲ):植株冠层光合作用数理模型

以1-2年生北加州黑核桃为试材,建立了具有较高分辨能力的植株群体结构、光分布模型和冠层光合作用模型．将植株冠层按叶面积指数划分为若干层次。上下层之间水平面上太阳辐照度按Monsi＆Saeki所提出的指数递减规律分布．冠层内太阳散射光的消光系数由冠层结构决定,而直射光的消光系数则决定于冠层结构与太阳在天空的位置．在同一层次。将叶片的叶倾角划分为6个等级。将叶片的水平位置划分为8个方位．设同一层次中水平面上的太阳辐照度相同。某一方位角和叶倾角的叶面的直接辐射由太阳视运动方程决定．以此为基础,分别计算“光斑区”和“遮荫区”内叶片的光合速率,并通过数值积分计算整个冠层的光合速率及光合日总量．用田间实测资料验证了冠层内太阳辐射分布模型和冠层光合作用模型．敏感性试验分析表明。模型对环境因子和生物学因素有良好的响应．     

Drought effects on canopy properties and productivity in thinned and unthinned Turkey oak stands

ABSTRACT

Drought responses, leaf area index (LAI), leaf characteristics and light extinction coefficient (k) were analysed in thinned and unthinned Turkey oak (Quercus cerris L.) stands at two sites: Valsavignone, in the Apennines, with a mild climate, and Caselli, near the Tyrrhenian coast, with a longer and more accentuated dry period in the summer. Turkey oak showed a good adaptability to drought due to a series of modifications in leaf characteristics, canopy properties and biomass allocation such as leaf area reduction, increased leaf thickness, smaller number of leaves and, at stand level, lower LAI, leaf biomass and LWR values and higher light extinction coefficients. In spite of the better environmental conditions and the higher LAI values, productivity was lower in the wet site. The differences in Turkey oak canopy properties, light extinction coefficients, LAI and their relations with drought and productivity are discussed.     

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.     

Carbon gain in a multispecies canopy: the role of specific leaf area and photosynthetic nitrogen-use efficiency in the tragedy of the commons

Models have been formulated for monospecific stands in which canopy photosynthesis is determined by the vertical distribution of leaf area, nitrogen and light. In such stands, resident plants can maximize canopy photosynthesis by distributing their nitrogen parallel to the light gradient, with high contents per unit leaf area at the top of the vegetation and low contents at the bottom. Using principles from game theory, we expanded these models by introducing a second species into the vegetation, with the same vertical distribution of biomass and nitrogen as the resident plants but with the ability to adjust its specific leaf area (SLA, leaf area:leaf mass). The rule of the game is that invaders replace the resident plants if they have a higher plant carbon gain than those of the resident plants. We showed that such invaders induce major changes in the vegetation. By increasing their SLA, invading plants could increase their light interception as well as their photosynthetic nitrogen-use efficiency (PNUE, the rate of photosynthesis per unit organic nitrogen). By comparison with stands in which canopy photosynthesis is maximized, those invaded by species of high SLA have the following characteristics: (1) the leaf area index is higher; (2) the vertical distribution of nitrogen is skewed less; (3) as a result of the supra-optimal leaf area index and the more uniform distribution of nitrogen, total canopy photosynthesis is lower. Thus, in dense canopies we face a classical tragedy of the commons: plants that have a strategy to maximize canopy carbon gain cannot compete with those that maximize their own carbon gain. However, because of this strategy, individual as well as total canopy carbon gain are eventually lower. We showed that it is an evolutionarily stable strategy to increase SLA up to the point where the PNUE of each leaf is maximized.     

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.     

播种量和施氮量对垄沟覆膜栽培冬小麦花后生理性状的影响

为协调冬小麦个体与群体间的关系,充分发挥旱作条件下垄沟栽培优势,以冬小麦品种小偃22为材料,采用二元二次正交旋转组合设计,通过田间试验研究了垄下集中施肥、垄上覆膜、膜际种植模式下播种量和施氮量对冬小麦花后生理性状的影响.结果表明： 花后叶面积指数、旗叶叶绿素含量和净光合速率均随施氮量的增加而增加.灌浆前中期叶面积指数随播种量的增加呈先增后稳的趋势;灌浆后期叶面积指数随播种量的增加而降低.随播种量的增加,旗叶的叶绿素含量和净光合速率降低,单株产量呈先减少后增加的趋势.适宜的播种量可以协调个体与群体间的矛盾,而适量增施氮肥有利于花后小麦生理性状的改善和产量的提高.在供试条件下,小偃22在播种量112.5 kg·hm-2与施氮量180～222 kg N·hm-2配置时,个体与群体的关系比较协调,花后叶面积指数较高,群体结构适宜,而且旗叶叶绿素含量、净光合速率和单茎产量较高,能获得高产.     

Structure of forest canopies as related to their primary productivity

Some structural features of forest canopy were analysed in relationto their role in photosynthetic production by forest communitieswhich are thought to produce more organic matter than herbaceouscommunities under the same environment. 1) Leaf area density was found to be much smaller in forestthan in herbaceous canopies. 2) Light extinction in forest canopy followed the BEER-LAMBERT'Slaw as was found for herbaceous canopy, though the coefficientof light extinction (K) was relatively small in the former. 3) A geometrical model was proposed to account for the smallvalue of K and the resultant large leaf area index, based onthe characteristically clustered distribution of tree leavesin forest canopy. 4) Stratification in forest community was interpreted as theuneven vertical distribution of leaf area density. 5) Deterioration of leaf functions, such as photosynthesis andrespiration, toward the bottom of forest canopy was noticed. 6) An attempt to estimate total canopy photosynthesis is presentedtaking this into consideration. 7) A new method for estimating total canopy respiration is proposedand discussed. ¹Contributions from JIBP-PT No. 36. The essential parts of thispaper were presented by KIRA to the IBP symposium The BiologicalBasis of Productivity held at Varna, Bulgaria, on April 4, 1968. (Received August 15, 1968; )     

播种量和施氮量对垄沟覆膜栽培冬小麦花后生理性状的影响

为协调冬小麦个体与群体间的关系,充分发挥旱作条件下垄沟栽培优势,以冬小麦品种小偃22为材料,采用二元二次正交旋转组合设计,通过田间试验研究了垄下集中施肥、垄上覆膜、膜际种植模式下播种量和施氮量对冬小麦花后生理性状的影响.结果表明:花后叶面积指数、旗叶叶绿素含量和净光合速率均随施氮量的增加而增加.灌浆前中期叶面积指数随播种量的增加呈先增后稳的趋势;灌浆后期叶面积指数随播种量的增加而降低.随播种量的增加,旗叶的叶绿素含量和净光合速率降低,单株产量呈先减少后增加的趋势.适宜的播种量可以协调个体与群体间的矛盾,而适量增施氮肥有利于花后小麦生理性状的改善和产量的提高.在供试条件下,小偃22在播种量112.5 kg hm-2与施氮量180 ～222 kg N·hm-2配置时,个体与群体的关系比较协调,花后叶面积指数较高,群体结构适宜,而且旗叶叶绿素含量、净光合速率和单茎产量较高,能获得高产.