Competition and Allometry in Kochia scoparia

Comparisons between crowded and uncrowded Kochia scoparia individualsdemonstrate pronounced effects of competition on plant allometryas well as on the distributions of different aspects of size.Non-destructive measurements of height and stem diameter and,for a subset of the populations, the number and length of leavesand branches, were taken at three times, and the plants wereharvested after the third measurement. The sequential measurementsafforded the opportunity to obtain information of the effectsof competition on allometric growth trajectories of individuals,as well as on static inter-individual allometric relationships. The distributions of most size measures appeared to be normalfor the uncrowded population. Crowded populations developeda negatively-skewed height distribution and a high-inequalitymass distribution, whereas the diameter distributions remainednormal. Plants grown without neighbours showed simple allometricrelationships between height, diameter and weight. For isolatedplants, the 'static' allometric relationship between plantsof different sizes and the allometric growth trajectory of individualswere similar. Crowded populations showed complex allometry;the static inter-individual relationships between height, diameterand weight were curvilinear (on log-log scale). There were largedifferences in the allometric growth slopes of uncrowded vs.crowded plants. Allometric relationships between stem diameterand plant mass, and between total length of leaves and totallength of branches, did not seem to be altered by competition. The data suggest that height was the most important aspect ofsize influencing future growth of individuals in the crowdedpopulation. Only plants above a certain height were able tocontinue to grow from the second to third measurement in thecrowded population. This supports the hypothesis that asymmetriccompetition for light is the cause of the allometric changesand of the increase in size variability due to competition.Copyright1994, 1999 Academic Press Allometric growth, allometry, competition, growth, Kochia     

Size structure of populations within populations: leaf number and size in crowded and uncrowded Impatiens pallida individuals

Summary We compared the size distributions of leaves on naturally-occurring crowded and experimentally thinned uncrowded individuals of Impatiens pallida in southeastern Pennsylvania. Crowding decreased the number of leaves on individual plants and altered the distribution of leaf size. Crowded individuals had smaller leaves, but the size (length) inequality of the leaf population did not change. The relationships between the height of a plant and the mean and maximum length of its leaves were significantly different for crowded and uncrowded plants. There were weak positive relationships between height and total leaf area, and height and total number of leaves for uncrowded plants, whereas crowded plants showed tighter but curvilinear relationships between these variables. Our results point out the strengths and the limitations of viewing canopies as populations of modules.     

Tree Size Inequality Reduces Forest Productivity: An Analysis Combining Inventory Data for Ten European Species and a Light Competition Model

Plant structural diversity is usually considered as beneficial for ecosystem functioning. For instance, numerous studies have reported positive species diversity-productivity relationships in plant communities. However, other aspects of structural diversity such as individual size inequality have been far less investigated. In forests, tree size inequality impacts directly tree growth and asymmetric competition, but consequences on forest productivity are still indeterminate. In addition, the effect of tree size inequality on productivity is likely to vary with species shade-tolerance, a key ecological characteristic controlling asymmetric competition and light resource acquisition. Using plot data from the French National Geographic Agency, we studied the response of stand productivity to size inequality for ten forest species differing in shade tolerance. We fitted a basal area stand production model that included abiotic factors, stand density, stand development stage and a tree size inequality index. Then, using a forest dynamics model we explored whether mechanisms of light interception and light use efficiency could explain the tree size inequality effect observed for three of the ten species studied. Size inequality negatively affected basal area increment for seven out of the ten species investigated. However, this effect was not related to the shade tolerance of these species. According to the model simulations, the negative tree size inequality effect could result both from reduced total stand light interception and reduced light use efficiency. Our results demonstrate that negative relationships between size inequality and productivity may be the rule in tree populations. The lack of effect of shade tolerance indicates compensatory mechanisms between effect on light availability and response to light availability. Such a pattern deserves further investigations for mixed forests where complementarity effects between species are involved. When studying the effect of structural diversity on ecosystem productivity, tree size inequality is a major facet that should be taken into account.     

The effects of density, spatial pattern, and competitive symmetry on size variation in simulated plant populations

Patterns of size inequality in crowded plant populations are often taken to be indicative of the degree of size asymmetry of competition, but recent research suggests that some of the patterns attributed to size-asymmetric competition could be due to spatial structure. To investigate the theoretical relationships between plant density, spatial pattern, and competitive size asymmetry in determining size variation in crowded plant populations, we developed a spatially explicit, individual-based plant competition model based on overlapping zones of influence. The zone of influence of each plant is modeled as a circle, growing in two dimensions, and is allometrically related to plant biomass. The area of the circle represents resources potentially available to the plant, and plants compete for resources in areas in which they overlap. The size asymmetry of competition is reflected in the rules for dividing up the overlapping areas. Theoretical plant populations were grown in random and in perfectly uniform spatial patterns at four densities under size-asymmetric and size-symmetric competition. Both spatial pattern and size asymmetry contributed to size variation, but their relative importance varied greatly over density and over time. Early in stand development, spatial pattern was more important than the symmetry of competition in determining the degree of size variation within the population, but after plants grew and competition intensified, the size asymmetry of competition became a much more important source of size variation. Size variability was slightly higher at higher densities when competition was symmetric and plants were distributed nonuniformly in space. In a uniform spatial pattern, size variation increased with density only when competition was size asymmetric. Our results suggest that when competition is size asymmetric and intense, it will be more important in generating size variation than is local variation in density. Our results and the available data are consistent with the hypothesis that high levels of size inequality commonly observed within crowded plant populations are largely due to size-asymmetric competition, not to variation in local density.     

Light regime in relation to plant population geometry

Plant population geometry effective in light utilization for photosynthesis was examined with the use of square-planted (SP) population models and the Monte Carlo technique. Varying SP populations were constructed by manipulating the structural variables, leaf area density, leaf size, leaf number, height/width ratio of unit stand and planting distance, of the unit stand with standard configurations treated in the second paper. Leaf area index was fixed to be 5, and the phyllotaxis, 1/3. The effects of these structural variables on the light extinction in the SP populations were made clear with light-beam emission experiments in a computer. Special combinations of the variables could make light extinction in the infinite population approximately linear with increasing leaf area index to obtain the highest photosynthesis of the foliage, i.e., each leaf layer from top to bottom of the population could uniformly utilize light energy for photosynthetic production.     

Effects of Ultraviolet-B Irradiance on Intraspecific Competition and Facilitation of Plants: Self-Thinning,Size Inequality,and Phenotypic Plasticity

(1) The effects of facilitation on the structure and dynamics of plant populations have not been studied so widely as competition. The UV-B radiation, as a typical environmental factor causing stress, may result in direct stress and facilitation. (2) The effects of UV-B radiation on intraspecific competition and facilitation were investigated based on the following three predictions on self-thinning, size inequality, and phenotypic plasticity: i) Self-thinning is the reduction in density that results from the increase in the mean biomass of individuals in crowded populations, and is driven by competition. In this study, the mortality rate of the population is predicted to decrease from UV-B irradiance. ii) The size inequality of a population increases with competition intensity because larger individuals receive a disproportionate share of resources, thereby leaving limited resources for smaller individuals. The second hypothesis assumes that direct stress decreases the size inequality of the population. iii) Phenotypic plasticity is the ability to alter one’s morphology in response to environmental changes. The third hypothesis assumes that certain morphological indices can change among the trade-offs between competition, facilitation, and stress. These predictions were tested by conducting a field pot experiment using mung beans, and were supported by the following results: (3) UV-B radiation increased the survival rate of the population at the end of self-thinning. However, this result was mainly due to direct stress rather than facilitation. (4) Just as competitor, facilitation was also asymmetric. It increased the size inequality of populations during self-thinning, whereas stress decreased the size inequality. (5) Direct stress and facilitation influence plants differently on various scales. Stress inhibited plant growth, whereas facilitation showed the opposite on an individual scale. Stress increased survival rate, whereas facilitation increased individual variability on the population scale. (6) Trade-offs between competitions, facilitation, and direct stress varied in different growing stages.     

Size-dependent growth and the development of inequality in maize,sunflower and soybean

Links were investigated between allometry of plant growth and dynamics of size structure of well-fertilized, irrigated crops of soybean (Glycine max L.), sunflower (Helianthus annuus L.) and maize (Zea mays L.) grown at standard plant-population densities (D), as in commercial crops (D = 30, 6 and 8.5 plants m-2, respectively), and at high densities (2D). Patterns of size-dependent growth of shoot and seed mass accumulation were distinctly different among species. In soybean and sunflower, non-linear relationships between size and subsequent growth led to strong hierarchical populations in terms of both shoot and seed biomass. Curvilinear (soybean) and sigmoid (sunflower) size-dependent growth determined strongly asymmetrical (soybean) and bimodal (sunflower) frequency distributions of shoot biomass indicating predominantly size asymmetrical competition among individuals. In comparison, a lower plant-to-plant variation coupled with a typical linear allometry of growth to plant size indicated symmetrical two-sided plant interference in maize. Despite the weak development of hierarchies in shoot biomass, a strong inequality in reproductive output developed in crowded populations of maize indicating an apparent breakage of reproductive allometry.     

Theoretical Relationships Between Mean Plant Size, Size Distribution and Self Thinning under One-sided Competition

As yet there is no comprehensive theory in plant populationecology to explain relationships between mean plant size, sizedistribution and self-thinning. In this paper, a new synthesisof plant monocultures is proposed. If the reciprocal relationshipbetween plant biomass and plant population density among variousstands of even-aged plant populations holds, the same reciprocalrelationship must exist between cumulative mass and cumulativenumber of plants from the largest individual within a population,assuming strict one-sided competition (which is an extreme conditionfor competition for light among plants). The two parametersof the relationship between cumulative mass and cumulative numberwithin a stand both correlate with maximum plant height in thestand. One parameter equals the reciprocal of the potentialmaximum plant mass per area, which is expressed by the productof maximum plant height and dry-matter density. The other parametercorrelates with the potential maximum individual plant mass,which is allometrically related to maximum plant height. Asa stand develops, the growth rate of the smallest individualswill become zero due to suppression from larger individuals,and they will die; i.e. self-thinning will occur. The slopeof the self-thinning line is expressed through the coefficientsof allometry between height and mass and between dry matterdensity and height. When the former coefficient is 3 and thelatter is 0, the gradient exactly corresponds to the value expectedfrom the 3/2 power rule, but it can take various values dependingon the values of the two coefficients. Competition among individualsdetermines size-density relationships among stands, which inturn determine the size structure of the stand. The size structureconstrains the growth of individuals and results in self-thinningwithin the stand.Copyright 1999 Annals of Botany Company. Monoculture, plant population, self-thinning, competition, hierarchy, size-structure.     

Old-growth red spruce forests as reservoirs of genetic diversity and reproductive fitness

Old-growth forests are assumed to be potential reservoirs of genetic diversity for the dominant tree species, yet there is little empirical evidence for this assumption. Our aim was to characterize the relationship of stand traits, such as age, height and stem diameter, with the genetic and reproductive status of old-growth and older second-growth stands of red spruce (Picea rubens Sarg.) in eastern Canada. We found strong relationships between height growth (a fitness trait) and measures of genetic diversity based on allozyme analyses in red spruce. The negative relationship between height and the proportion of rare alleles suggests that high proportions of these rare alleles may be deleterious to growth performance. Latent genetic potential, however, showed a significant and positive relationship with height. Stand age was not correlated to height, but was correlated to seedling progeny height. In late-successional species such as red spruce, age and size (e.g., height and stem diameter) relationships may be strongly influenced by local stand disturbance dynamics that determine availability of light, growing space, moisture and nutrients. In larger and older stands, age appeared to provide a good surrogate measure or indicator for genetic diversity and progeny height growth. However, in smaller and more isolated populations, these age and fitness relationships may be strongly influenced by the effects of inbreeding and genetic drift. Therefore, older populations or old-growth forests may represent superior seed sources, but only if they are also of sufficient size and structure (e.g., stem density and spatial family structure) to avoid the effects of inbreeding and genetic drift. Thus, larger and older forests appear to have an important evolutionary role as reservoirs of both genetic diversity and reproductive fitness. Given the rapid environmental changes anticipated (as a result of climate change, increasing population isolation through fragmentation, or following the introduction of exotic pests and diseases) these older populations of trees may have a valuable function in maintaining the adaptive potential of tree species.     

Modeling the growth of individuals in crowded plant populations

Aims We present an improved model for the growth of individuals in plant populations experiencing competition.Methods Individuals grow sigmoidally according to the Birch model, which is similar to the more commonly used Richards model, but has the advantage that initial plant growth is always exponential. The individual plant growth models are coupled so that there is a maximum total biomass for the population. The effects of size-asymmetric competition are modeled with a parameter that reflects the size advantage that larger individual have over smaller individuals. We fit the model to data on individual growth in crowded populations of Chenopodium album .Important findings When individual plant growth curves were not coupled, there was a negative or no correlation between initial growth rate and final size, suggesting that competitive interactions were more important in determining final plant size than were plants' initial growth rates. The coupled growth equations fit the data better than individual, uncoupled growth models, even though the number of estimated parameters in the coupled competitive growth model was far fewer, indicating the importance of modeling competition and the degree of size-asymmetric growth explicitly. A quantitative understanding of stand development in terms of the growth of individuals, as altered by competition, is within reach.     

尾叶桉人工林种群密度的研究

探讨了5.6年生尾叶桉种群密度与冠幅、胸径、树高、立木单株材积、林分蓄积量、木材性质及保存率等的作用规律和相关模型.结果表明,密度对胸径、立木单株材积、冠幅及枝下高的影响达到极显著水平;对蓄积量、木材纤维宽度的影响达显著水平;对树高、木材气干密度和木材纤维长度虽有一定的影响,但不显著.其中,密度与蓄积量、枝下高、木材纤维宽度呈正相关关系;而与胸径、立木单株材积、冠幅呈负相关关系.此外,尾叶桉具有较宽的合理密度范围;作为短周期浆纸林,其最佳密度应确定为2000株·hm_-2.     

尾叶桉人工林种群密度的研究

探讨了５．６年生尾叶桉种群密度与冠幅、胸径、树高、立木单株材积、林分蓄积量、木材性质及保存率等的作用规律和相关模型。结果表明,密度对胸径、立木单株材积、冠幅及枝下高的影响达到极显著水平;对蓄积量、木材纤维宽度的影响达显著水平;对树高、木材气干密度和木材纤维长度虽有一定的影响,但不显著。其中,密度与蓄积量、枝下高、木材纤维宽度呈正相关关系;而与胸径、立林单株材积、冠幅呈负相关关系。此外,尾叶桉具有较     

Tree growth and competition in a Betula platyphylla-Larix cajanderi post-fire forest in central Kamchatka

BACKGROUND AND AIMS: Fire is the dominant disturbance in central Kamchatka boreal forests, yet patterns and mechanisms of stand recovery have not been investigated. METHODS: Measurements were made of 1433 stems > or =1.3 m height and annual radial increments of 225 randomly selected trees in a 0.4-ha plot of a 53-year-old fire-origin mixed-species stand to examine the spatio-temporal variation in establishment, growth, size inequality and the mode of competition among individual trees. Growth variations were related to tree size, age and local interference with neighbours. KEY RESULTS: Betula platyphylla formed the main canopy following a fire in 1947, with Larix cajanderi and Pinus pumila progressively reinvading the lower tree and shrub stratum. Most B. platyphylla originated from sprouts in small patches (polycormons) during the first 15 post-fire years. Betula platyphylla had normal distributions of diameter and age classes, but negatively skewed height distribution, as expected from shade-intolerant, pioneer species. Larix cajanderi had fewer tall and many short individuals. The smaller and younger B. platyphylla grew disproportionately more in diameter than larger trees from 1950 to 1975, and hence stem size inequalities decreased. The reverse trend was observed from 1995 to 2000: larger trees grew more, indicating an increasing asymmetry of competition for light. Betula platyphylla had steady diameter growth in the first 25 post-fire years, after which the growth declined in smaller trees. Neighbourhood analysis showed that the decline resulted from increased competition from taller neighbours. CONCLUSIONS: The observed growth patterns suggest that mode of interactions altered during stand development from early stages of weak competition for soil resources released by fire to later stages of asymmetric competition for light. Asymmetric crown competition started later than reported in other studies, which can be attributed to the lower stem density leaving much space for individual growth, greater relative importance of below-ground competition in this site of nutrient-poor volcanic soil, and the vegetative origin of B. platyphylla. Larix cajanderi growing under B. platyphylla had steady diameter growth during the first 20 years, after which growth declined. It is suggested that early succession fits the tolerance model of succession, while inhibition dominates in later stages.     

SEASONAL CHANGES IN SIZE STRUCTURE OF SARGASSUM AND TURBINARIA POPULATIONS (PHAEOPHYCEAE) ON TROPICAL REEF FLATS IN THE SOUTHERN RED SEA1

Seasonal variation in density, thallus length and biomass, population size structure, and allometric length‐biomass relationships was investigated in populations of Sargassum ilicifolium (Turner) C. Agardh, Sargassum subrepandum (Forssk.) C. Agardh, and Turbinaria triquetra (J. Agardh) Kütz. (Phaeophyceae) on shallow reef flats in the southern Red Sea. Thallus length and biomass varied strongly with season, with the highest values occurring in the cooler months. Thallus densities showed no significant temporal variation. Log‐total biomass versus log‐density relationships were positive throughout the growth season without any decrease in the slope of the relationship. In two populations, biomass‐density combinations approached the interspecific biomass‐density line, but the massive annual shedding of modules occurred before self‐thinning would set in. Allometric length‐biomass relationships varied with season in all populations and were associated with seasonal module initiation, growth, and shedding. Evidence of a strong asymmetric competition was found in two high‐density populations. These populations showed a predominance of small thalli during peak development, asymmetrical Lorenz curves, increasing Gini coefficients, and increasing thallus length relative to biomass during the main growth phase. In two other less crowded populations, small thalli were absent during peak development, Lorenz curves were symmetrical, and Gini coefficients decreased during the main growth phase. In these populations, size equalization appears to be due to responses at the modular level rather than size‐dependent mortality. We conclude that changes in size structure in this highly seasonal environment are determined by module dynamics, modified by asymmetric competition in some populations, with a minor role of recruitment and no regulatory effect of self‐thinning.     

The Processes of Height-rank Determination Among Individuals and Neighbourhood Effects inChenopodium albumL. Stands

The height ofChenopodium albumL. plants grown in monocultureat three different densities was followed throughout the growingseason to examine size-rank determination processes with specialreference to the effects of neighbourhood conditions. Changesin height rank of plants in the stands were assessed by therank correlation between final height and the height at eachmeasurement during the growing season. The height ranks of plantswere almost fixed 1–2 weeks after canopy closure whenthe stand height was 10–20% of final stand height, andfixation occurred earlier in the denser plot. At each measurement,the effects of neighbourhood were evaluated as the partial correlationcoefficient between height growth and neighbourhood index withheight held constant (r_GN.H), in which competitive asymmetrywas incorporated. During the early period of the growing season,r_GN.Hwasnon-significant or positive (plants with taller and/or closerneighbours elongated faster), indicating no local competition.Just after canopy closure,r_GN.Hbecame negative, indicating localcompetition. A plant's rank changed only in an initial shortperiod of the competition. Plants occupying the upper canopyof stands at the end of the growing season were distinguishedby greater height growth during the initial short period ofcompetition after canopy closure, although these plants werenot necessarily taller before the onset of local competition.These results suggest that the fate of a plant in a crowdedstand is determined in the early stage of stand development.Copyright1999 Annals of Botany Company Height growth, neighbourhood competition, local competition, height-rank of plants in population, size difference, asymmetric competition,Chenopodium albumL.     

Stem extension and mechanical stability of Xanthium canadense grown in an open or in a dense stand

Background and Aims

Plants in open, uncrowded habitats typically have relatively short stems with many branches, whereas plants in crowded habitats grow taller and more slender at the expense of mechanical stability. There seems to be a trade-off between height growth and mechanical stability, and this study addresses how stand density influences stem extension and consequently plant safety margins against mechanical failure.

Methods

Xanthium canadense plants were grown either solitarily (S-plants) or in a dense stand (D-plants) until flowering. Internode dimensions and mechanical properties were measured at the metamer level, and the critical buckling height beyond which the plant elastically buckles under its own weight and the maximum lateral wind force the plant can withstand were calculated.

Key Results

Internodes were longer in D- than S-plants, but basal diameter did not differ significantly. Relative growth rates of internode length and diameter were negatively correlated to the volumetric solid fraction of the internode. Internode dry mass density was higher in S- than D-plants. Young''s modulus of elasticity and the breaking stress were higher in lower metamers, and in D- than in S-plants. Within a stand, however, both moduli were positively related to dry mass density. The buckling safety factor, a ratio of critical buckling height to actual height, was higher in S- than in D-plants. D-plants were found to be approaching the limiting value 1. Lateral wind force resistance was higher in S- than in D-plants, and increased with growth in S-plants.

Conclusions

Critical buckling height increased with height growth due mainly to an increase in stem stiffness and diameter and a reduction in crown/stem mass ratio. Lateral wind force resistance was enhanced due to increased tissue strength and diameter. The increase in tissue stiffness and strength with height growth plays a crucial role in maintaining a safety margin against mechanical failure in herbaceous species that lack the capacity for secondary growth.     

Inequality of size and size increment in Pinus banksiana in relation to stand dynamics and annual growth rate

BACKGROUND AND AIMS: Changes in size inequality in tree populations are often attributed to changes in the mode of competition over time. The mode of competition may also fluctuate annually in response to variation in growing conditions. Factors causing growth rate to vary can also influence competition processes, and thus influence how size hierarchies develop. METHODS: Detailed data obtained by tree-ring reconstruction were used to study annual changes in size and size increment inequality in several even-aged, fire-origin jack pine (Pinus banksiana) stands in the boreal shield and boreal plains ecozones in Saskatchewan and Manitoba, Canada, by using the Gini and Lorenz asymmetry coefficients. KEY RESULTS: The inequality of size was related to variables reflecting long-term stand dynamics (e.g. stand density, mean tree size and average competition, as quantified using a distance-weighted absolute size index). The inequality of size increment was greater and more variable than the inequality of size. Inequality of size increment was significantly related to annual growth rate at the stand level, and was higher when growth rate was low. Inequality of size increment was usually due primarily to large numbers of trees with low growth rates, except during years with low growth rate when it was often due to small numbers of trees with high growth rates. The amount of competition to which individual trees were subject was not strongly related to the inequality of size increment. CONCLUSIONS: Differences in growth rate among trees during years of poor growth may form the basis for development of size hierarchies on which asymmetric competition can act. A complete understanding of the dynamics of these forests requires further evaluation of the way in which factors that influence variation in annual growth rate also affect the mode of competition and the development of size hierarchies.     

Not only light quality but also mechanical stimuli are involved in height convergence in crowded Chenopodium album stands

? In crowded stands, height is often similar among dominant plants, as plants adjust their height to that of their neighbours (height convergence). We investigated which of the factors, light quality, light quantity and mechanical stimuli, is primarily responsible for stem elongation and height convergence in crowded stands. ? We established stands of potted Chenopodium album plants. In one stand, target plants were surrounded by artificial plants that were painted black to ensure that the light quality was not modified by their neighbours. In a second stand, target plants were surrounded by real plants. In both stands, one-half of the target plants were anchored to stakes to prevent flexing by wind. The target plants were lifted or lowered by 10?cm to test whether height convergence was affected by the different treatments. ? Stem length was affected by being surrounded by artificial plants, anchoring and pot elevation, indicating that light quality, light quantity and mechanical stimuli all influenced stem elongation. Height convergence did not occur in the stand with artificial plants or in anchored plants. ? We conclude that light quality and mechanical stimuli are important factors for the regulation of stem growth and height convergence in crowded stands.     

Patterns of Trunk Diameter, Tree Height and Crown Depth in Crowded Abies Stands

The size structure of trees in crowded, even-aged Abies (fir)stands of ‘Shimagare’ or ‘wave-regenerated’sub-alpine forests is analyzed. Tree-height distributions showconsistently smaller variation and less positive skewness thanthe distributions of trunk diameter and crown depth (tree heightminus height of the lowest branch). This difference is associatedwith changes in the relationships between trunk diameter, treeheight and crown depth as stands age. These, in turn, resultfrom self-pruning of the lower foliage crown due to competitionfor light in crowded stands. Abies, diameter-height curve, competition, size distribution, stand development, tree geometry, wave-regeneration