Biomass Allocation and Light Partitioning Among Dominant and Subordinate Individuals inXanthium canadenseStands

Patterns of above-ground biomass allocation and light captureby plants growing in dense stands or in isolation were studiedin relation to their height. A canopy model was developed tocalculate light absorption by individual plants. This modelwas combined with data on canopy structure and patterns of biomassallocation for solitary plants and for plants of different heightsin dense mono-specific stands of the dicotyledonous annualXanthiumcanadenseMill. There were four stands, and stand height increasedwith age and nutrient availability. The allometric relationshipbetween height and mass differed considerably between plantsin stands and those growing in isolation and also between plantsof different heights within stands. The proportion of shootmass in leaf laminae (LMR) decreased with increasing plant height,but solitary plants had a higher LMR than competing plants ofthe same height. Thus, in contrast to previous assumptions,LMR of competing plants is not strictly determined by biomechanicalconstraints but results from a plastic shift in biomass allocationin response to competition. Average leaf area per unit leafmass (SLA) decreased with increasing photosynthetic photon fluxdensity (PPFD) independent of nutrient availability. Consequently,taller, more dominant plants in stands had a lower leaf arearatio (LAR: LAR=LMRxSLA) than shorter, more subordinate plants.Dominant plants absorbed more light both per unit leaf area(     

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.     

Canopy structure and nitrogen distribution in dominant and subordinate plants in a dense stand of Amaranthus dubius L. with a size hierarchy of individuals

The objective was to investigate how nitrogen allocation patterns in plants are affected by their vertical position in the vegetation (i.e. being either dominant or subordinate). A garden experiment was carried out with Amaranthus dubius L., grown from seed, in dense stands in which a size hierarchy of nearly equally aged individuals had developed. A small number of dominant plants had most of their leaf area in the highest layers of the canopy while a larger number of subordinate plants grew in the shade of their dominant neighbours. Canopy structure, vertical patterns of leaf nitrogen distribution and leaf photosynthetic characteristics were determined in both dominant and subordinate plants. The light distribution in the stands was also measured. Average N contents per unit leaf area (total canopy nitrogen divided by the total leaf area) were higher in the dominant than in the subordinate plants and this was explained by the higher average MPA (leaf dry mass per unit area) of the dominant plants. However, when expressed on a weight basis, average N contents (LNC_av; total canopy N divided by the total dry weight of leaves) were higher in the subordinate plants. It is possible that these higher LNC_av values reflect an imbalance between carbon and nitrogen assimilation with N uptake exceeding its metabolic requirement. Leaf N content per unit area decreased more strongly with decreasing relative photon flux density in the dominant than in the subordinate plants showing that this distribution pattern can be different for plants which occupy different positions in the light gradient in the canopy. The amount of N which is reallocated from the oldest to the younger, more illuminated leaves higher up in the vegetation may depend on the sink strength of the younger leaves for nitrogen. In the subordinate plants, constrained photosynthetic activity caused by shading might have reduced the sink intensity of these leaves.     

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.     

Light partitioning among species and species replacement in early successional grasslands

Abstract. We studied canopy structure, shoot architecture and light harvesting efficiencies of the species (photon flux captured per unit above‐ground plant mass) in a series of exclosures of different age (up to 4.5 yr) in originally heavily grazed grassland in N Japan.Vegetation height and Leaf Area Index (LAI) increased in the series and Zoysia japonica, the dominant in the beginning, was replaced by the much taller Miscanthus sinensis. We showed how this displacement in dominance can be explained by inherent constraints on the above‐ground architecture of these two species. In all stands light capture of plants increased with their above‐ground biomass but taller species were not necessarily more efficient in light harvesting. Some subordinate species grew disproportionally large leaf areas and persisted in the shady undergrowth. Some other species first grew taller and managed to stay in the better‐lit parts of the canopy, but ultimately failed to match the height growth of their neighbours in this early successional series. Their light harvesting efficiencies declined and this probably led to their exclusion. By contrast, species that maintained their position high in the canopy managed to persist in the vegetation despite their relatively low light harvesting efficiencies. In the tallest stands ‘later successional’ species had higher light harvesting efficiencies for the same plant height than ‘early successional’ species which was mostly the result of the greater area to mass ratio (specific leaf area, SLA) of their leaves. This shows how plant stature, plasticity in above‐ground biomass partitioning, and architectural constraints determine the ability of plants to efficiently capture light, which helps to explain species replacement in this early successional series.     

Leaf nitrogen distribution and whole canopy photosynthetic carbon gain in herbaceous stands

The amount of photosynthetically-active photon flux density incident upon a leaf and the nitrogen content of that leaf strongly affect the photosynthetic carbon gain of that leaf. Therefore, the canopy structure of a stand, affecting the light climate in the canopy, and the leaf nitrogen distribution pattern in the canopy, affect the carbon gain of the whole canopy. This review discusses the results of studies directed to this problem and obtained so far in open and in dense canopies of stands of herbaceous, monocotyledonous or dicotyledonous, plants in their growing or flowering stages. It is found that the leaf nitrogen distribution pattern in the canopy is vertically non-uniform, and in dense stands more strongly so than in open stands. The leaf nitrogen distribution pattern in most canopies closely approaches an optimal pattern in that it maximizes whole canopy potential carbon gain as calculated for the actual total leaf nitrogen content and leaf area index of the stand. The resulting increase in potential carbon gain as compared to a uniform leaf nitrogen distribution pattern is considerable and it is larger in dense stands than in open stands. For at least some dense stands simulation studies show that with the available total leaf nitrogen content, whole canopy carbon gains could still be considerable higher had a lower leaf area index been developed.     

Shoot dynamics of the giant grass Gynerium sagittatum in Peruvian Amazon floodplains,a clonal plant that does show self-thinning

The giant rhizomatous grass Gynerium sagittatum is an early successional species that forms dense monocultures in Peruvian Amazon floodplains. We studied the shoot population structures by recording shoot densities and shoot heights. Leaf areas and stem volumes were allometrically estimated. Stands of two varieties of G. sagittatum were examined that differ in height and in the degree of shoot branching. In stands of increasing age, marked decreases in shoot densities were accompanied with an increase in mean shoot size. Self-thinning was indicated by the negative correlation between log stem volume per unit ground area and log shoot density, significant at least for one of the two varieties. The difference in thinning slope between the varieties could be largely accounted for by their different shoot geometry, as was revealed by calculations based on the allometric model of Weller (1987b). The relationship between log leaf area per shoot and log shoot density was significantly negative with slopes close to –1. Shoot size inequalities decreased with increasing mean stem volume per shoot, probably as a result of density-dependent mortality of the smaller shoots. All of these results accord with expectations for shoot self-thining. Gynerium sagittatum is the first clear example of a clonal plant species that exhibits self-thining in natural monospecific stands. It is argued that self-thinning occurs in this giant tropical grass because its shoots are perennial and do not experience seasonal die-back (periodic density-independent mortality), in contrast to many of the clonal plant species that have been studies so far.     

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

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

Shoot dynamics and architecture of saplings in Fagus crenata across its geographical range

 Relationships between leaf or shoot size, number, and arrangement in response to light were investigated to test the hypothesis that these characteristics are linked. In order to test this hypothesis, the divergence in allometry and shoot dynamics in saplings of Japanese beech (Fagus crenata) obtained from four populations and having different leaf sizes were examined in a nursery under both full sun and shade conditions. Trees with different leaf sizes also showed large differences in canopy structure, particularly when shade-grown saplings were compared. The final leaf mass distributions of the large-leaf populations were conical or “bottom - heavy”, while those of the small-leaf populations were planar or “top - heavy”. The slope of the allometric relations between leaf mass and shoot and branch mass in small-leaved populations were steeper than those in large-leaf populations. The four populations were classified into two growth types: populations producing a few large leaf and shoot modules corresponded to “stem growth type”, and those producing many small leaf and shoot modules corresponded to “leaf growth type”. These kinds of intra-specific variation in architecture and growth of F. crenata trees may influence the structure and dynamics of forests in accordance with differences in competitive ability or sensitivity to disturbances such as windstorm. Received: 18 March 1997 / Accepted: 21 October 1997     

陆生生境中喜旱莲子草的生长模式

喜旱莲子草(Alternanthera philoxeroides)原产南美洲, 后被引入到北美洲、大洋洲、东南亚和中国等地, 成为一个世界性的外来入侵种。对喜旱莲子草陆生种群的有效控制一直是一个难题。本文中通过种植实验建立了陆生生境中喜旱莲子草主枝长、生物量、叶面积和斑块面积等的生长模型。结果表明: (1)喜旱莲子草的主枝长、生物量、叶面积和斑块面积等均表现为指数式生长, 其日增长率(%)分别为4.28、11.27、11.59和8.67。(2)喜旱莲子草的地上重(x)－地下根茎重(y)的异速生长指数b约为3/4(01), 即总重和叶面积相对于主枝长呈二次幂增长, 由此可进一步推出总重和叶面积与斑块面积成正比; 生物量－叶面积的异速生长指数b约为1, 为等速生长(b＝1), 即单位生物量所支持的叶面积不随植株大小的变化而变化(冠层恒定性)。其叶面积比为88.24 cm²/g, 比叶面积为287.97 cm²/g。通过本研究期望对喜旱莲子草陆生局域斑块的生长进行预测, 同时为进一步建立其控制模型提供基础数据, 为制定经济有效的控制对策提供科学依据。     

Mass loss,fungal colonisation and nutrient dynamics of Phragmites australis leaves during senescence and early aerial decay

This study examined the mass loss, fungal biomass, and nutrient dynamics of standing Phragmites australis leaf blades during senescence and early decay in littoral reed stands of two hardwater lakes. Green living leaves were tagged at defined canopy heights in early autumn (late August or early September) and periodically collected until all leaf blades had fallen off the parent shoot. Samples were analysed for leaf dry mass remaining, fungal biomass associated with leaves (ergosterol concentrations), and nitrogen and phosphorus concentrations. Considerable mass loss of leaves occurred in the standing position (up to 28%). Nitrogen and phosphorus concentrations of leaves decreased substantially with time (by 39–77%), indicating that a major portion of these nutrients was translocated to the rhizome during senescence. Fungal biomass associated with leaves increased during the study period, reaching an estimated maximum of about 40 mg g⁻¹ of leaf dry mass. Fungal biomass was negatively correlated with leaf N and P concentrations. The observed patterns of leaf mass loss, nutrient dynamics, and fungal biomass were consistent with the successive senescence and death of leaves from the shoot base to its tip. The results of this study point to a notable mass loss of P. australis leaf blades in the standing position, which appears to be mediated by both plant and microbial processes. Nutrient dynamics, in contrast, appear to be largely governed by plant processes.     

Effect of 3D nitrogen, dry mass per area and local irradiance on canopy photosynthesis within leaves of contrasted heterogeneous maize crops

BACKGROUND AND AIMS: Nitrogen partitioning within stands has been described fairly comprehensively, especially for C(3) plants in dense stands where the horizontal heterogeneity of foliage distribution is relatively small. Nitrogen has been shown to be distributed vertically and in parallel to light, maximizing carbon assimilation and stand productivity. Conversely, row crops such as maize (C(4) plants) are characterized by strong horizontal heterogeneity of foliage distribution, and a three-dimensional (3D) approach is required to investigate the combined effect of spatial distribution of nitrogen and light on canopy photosynthesis. MODEL: The 3D geometry of maize canopies was modelled with varying densities and at different developmental stages using plant digitizing under field conditions. For lamina parts, photosynthesis was measured and nitrogen content per unit area (N(a)) was described from analysis of nitrogen content per unit mass (N(m)) and dry mass per unit area (M(a)). Hyperbolic relationships between photosynthesis at irradiance saturation (P(max)) and N(a) were established as well as a linear relationship between dark respiration (R(d)) and N(a), whereas quantum efficiency (alpha) was found to be independent of N(a). KEY RESULTS AND CONCLUSIONS: N(m), M(a) and N(a) were shown to change over time vertically (i.e. between laminae), which has been largely reported previously, and horizontally (i.e. within laminae), which has scarcely been described previously. Even if M(a) played a major role in N(a), a strong relationship between N(a) and M(a) could not be demonstrated, whereas several previous studies have found that N(a) was essentially related to M(a) rather than N(m). From simulations of radiative exchange using a 3D volume-based approach and lamina photosynthesis using a hyperbola, it was shown that real patterns of N(a) partitioning could increase daily crop photosynthesis by up to 8 % compared with uniform patterns of N(a), especially for the earliest stages of stand development.     

Effects of geminivirus infection and growth irradiance on the vegetative growth and photosynthetic production of Eupatorium makinoi

Eupatorium makinoi plants with or without geminivirus infection were grown in shading frames with 70, 15 and 5.5% sunlight. Growth characteristics of these plants in the early vegetative phase were compared by means of growth analysis. We also measured leaf photosynthetic gas exchange rates and examined relationships between leaf photosynthesis and whole-plant growth. Relative growth rate (RGR=(1/W)×(dW/dt), where W is plant dry mass) of virus-infected plants was lower than that of uninfected plants under all three light conditions. The reduction of RGR by infection was increased with irradiance. The net assimilation rate (NAR=(1/A)×(dW/dt), where A is total leaf area of the plant) was also reduced both by infection and shading. NARs that were estimated from light-response curves of leaf photosynthesis, in situ measurements of irradiance, and respiration rates of leaves, stems and below-ground parts, agreed very well with the values obtained by conventional growth analysis techniques. Decreases in the estimated NAR value from infection and shading were mostly explained by the decreases in leaf photosynthesis. These results clearly showed that lowered RGR in virus-infected plants was attributed mainly to impaired photosynthesis in virus-infected leaves.     

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.     

Do bryophyte shoot systems function like vascular plant leaves or canopies? Functional trait relationships in Sphagnum mosses (Sphagnaceae)

Vascular plant leaf traits that influence photosynthetic function form the basis of mechanistic models of carbon exchange. Given their unique tissue organization, bryophytes may not express similar patterns. We investigated relationships among tissue, shoot, and canopy traits, and their associations with photosynthetic characteristics in 10 Sphagnum species. Trait relationships were organized around a primary dimension accounting for 43% of variation in 12 traits. There was no significant relationship between nitrogen content of shoot systems and maximum photosynthesis expressed on mass (A(mass)) or area (A(area)) bases due to nitrogen sequestration and storage within the canopy interior. This pattern differs from the distribution of nitrogen in vascular plant canopies. Thus, nitrogen and its relationship to carbon uptake in Sphagnum shoots does not conform to patterns of either vascular plant leaves or canopies. Species that concentrate biomass and nitrogen in the capitulum have enhanced rates of A(mass) and A(area). Consequently, A(area) was positively associated with N(area) of the capitulum only. Overall, water content and carotenoid concentration were the strongest predictors of both A(mass) and A(area) and these were expressed as inverse relationships. The relationships of plant traits in Sphagnum defines a principal trade-off between species that tolerate environmental stress and those that maximize carbon assimilation.     

Self-thinning lines differ with fertility level

The effect of variations in fertility level of the substrate on the self-thinning lines followed by populations of Ocimum basilicum L. was investigated experimentally by establishing populations over a range of densities at two fertility levels. Populations from each fertility level followed different self-thinning lines for shoot biomass. Self-thinning began at a lower biomass in populations grown at the higher fertility level; the subsequent slope of the thinning line was –0.5 for these stands on a log shoot biomass versus log density plot. The slope of the self-thinning line was flatter (–0.29) at the lower fertility level. Fitting the self-thinning line by the Structural Relationship rather than the Major Axis made little difference to line estimates. Biomass packing differed with fertility level, with plants from the higher fertility stands requiring more canopy volume for given shoot biomass than plants from lower fertility levels. Biologically, this would mean shoot competition intensified more rapidly at the higher fertility level as biomass accumulated in stands. The difference in slope between fertility levels was associated with changes above- and belowground. The radial extension of the canopy versus shoot mass relationships of individual plants differed with fertility level. Plants at the lower fertility level allocated more biomass to root growth, and had less leaf area per unit root length. The differences in slope of the self-thinning lines may have been because of differences in the radial extension of the canopy versus shoot mass relationships of individual plants at each fertility level, and/or to an increase in root competition at the lower fertility level.     

Vertical leaf nitrogen distribution in relation to nitrogen status in grassland plants

Vertical gradients of leaf nitrogen (N) per unit leaf area (NLA) are viewed as plastic responses that optimize N utilization with respect to carbon assimilation. However, it has been shown that plant species, sowing density and N availability affect the steepness of the NLA gradient relative to the photon flux density (PFD) gradient. This paper tests the hypothesis that such variation is related to the N status of the plant. The N status was analysed using the concept of the critical N concentration (Ncrit) in which shoot N per unit dry mass (NSM) decreases with shoot mass, and a negative deviation of actual NSM from Ncrit indicates N shortage in the plant. The hypothesis was tested with contrasting grassland species Medicago sativa, Dactylis glomerata and Taraxacum officinale by varying PFD and N availability, plant density and hierarchical positions of individuals within stands. Combinations of all treatments showed a general negative correlation between the N allocation coefficient (i.e. the slope of the NLA-PFD relationship) and NSM for all three species. Thus, NLA, relative to PFD, gradients became steeper with increasing shoot mass and increasing N shortage in the plant. These data are consistent with the view that internal N availability is an important factor in modifying the NLA gradient.     

千烟洲针叶林的比叶面积及叶面积指数

根据实测数据计算了湿地松(Pinus elliotii)、马尾松(P. massoniana) 和杉木(Cunninghamia lanceolata)不同年龄、不同类型叶片的生物量和比叶面积,并结合样地调查数据和相对生长方程计算了中国科学院千烟洲试验站20年生湿地松林、马尾松林、杉木林和针叶混交林的叶面积指数。根据拟合结果,选择如下方程计算3个树种的叶生物量:湿地松W=12.074 1D^{2.151 5}、马尾松W=6.972 7D^{2.197 3}和杉木W=5.261 9D^{2.302 7}。湿地松林的叶生物量(0.822 kg·m^-2)最大,其次为针叶混交林(0.679 kg·m^-2),马尾松林和杉木林相差不大(分别为林0.528和0.572 kg·m^-2)。不同树种、不同年龄、不同类型叶片的比叶面积比较发现,新叶的比叶面积大于老叶,三针一束叶的比叶面积略大于两针一束叶,马尾松的平均半比表面积(8.62 m²·kg^-1)大于湿地松(6.04 m²·kg^-1)和杉木(7.91 m²·kg^-1)。胸径与单木叶片半表面积之间的经验方程为:湿地松LA=0.073D^{2.151 5}、马尾松LA=0.060D^{2.197 3}和杉木LA=0.042D^{2.302 7}。据此计算湿地松林的叶面积指数为5.03,马尾松林和杉木林为4.31,针叶混交林为4.77,该结果比利用CI-110植被冠层数字图像仪测得的结果偏大。     

Survivorship,growth and self-thinning in Banksia ericifolia

High-density (dense) and low-density (sparse) plots were set up in naturally sown monospecific stands of Banksia ericifolia in coastal heath, 3 years after fire. This was done both in high-growth and low-growth areas. Plant mortality was recorded quarterly, and two harvests were made at 6 and 9 years to sample growth. Density-independent mortality at an exponential rate was observed in the low-growth treatments at both densities, and in the high-growth sparse treatment. Growth level affected mortality, with the half-life of populations in the high-growth sparse plots being double that of populations in the low-growth plots. Density-dependent mortality (self-thinning) was seen only in the high-growth dense plots. Seasonal effects on mortality were slight; maximum mortality was observed in the spring-summer period in plots subject to density-independent mortality, and in the winter-spring quarter in plots that had self-thinned. Yields in the high-growth plots and the low-growth dense plots were high for heath vegetation. The self-thinning populations did not exceed White's (1985) upper boundary for thinning lines of log intercept (K) = 5 on standardized axes. The data suggested a log intercept value in the range 4.8–4.9 in the high-growth stands assuming a thinning-line slope of – 1.5. Banksia ericifolia (a large shrub/small tree) has a high mean plant weight per given thinning density compared with trees, where an upper limit of log K= .4 has been suggested by White (1985). The volume of canopy space per plant in B. ericifolia is not unusual compared with other species. The amount of biomass packed into a given volume of canopy space was high in this Banksia, achieved by having leaves with a low ratio of area to weight (specific leaf area, SLA). For given values of density, leaf area index and proportion of shoot as leaf, plants with a low SLA will be several times heavier than plants with a high SLA. This achieves a high biomass to volume ratio without an erectophile canopy and may explain the high intercept seen for thinning lines of conifers.     

Seasonal dynamics of leaf C,N and P stoichiometry in plants of typical steppe in Nei Mongol,China

分析植物叶片(C)、氮(N)、磷(P)含量及其比值的季节动态, 不仅有助于认识植物生长发育和养分吸收利用等生理生态过程, 也有利于认识植物化学计量的动态平衡关系。该文选择内蒙古典型温带草原18种常见植物, 在生长季的6-9月, 每半月一次进行连续采样, 在此基础上分析了叶片C、N、P含量及其比值在生长季内的变化。主要结果: 1)植物叶片C、N、P含量及其比值的季节性变化在不同功能类群间不同步, 其中叶片N、P含量的季节变化体现了明显的稀释作用。2)叶片C、N、P含量及其比值在不同功能类群间差异显著, 单子叶、多年生禾草类的叶片N、P含量显著低于双子叶和多年生杂类草植物, 而其叶片C:N、C:P则高于双子叶和多年生杂类草植物。3)叶片N、P含量显著正相关, 叶片C:N和C:P分别与N和P含量显著负相关, 可能体现了植物体内营养元素间的内在耦合机制。4)叶片N含量与C:N, 叶片P含量与C:P以及叶片N含量与P含量均呈现等速生长关系, 且等速生长关系在生长季保持稳定。