Crowding in clonal seaweeds: Does self-thinning occur in Mastocarpus papillatus shortly before stand biomass peaks?

Fronds from crowded stands of clonal seaweeds, particularly those in which holdfasts are mostly perennial and are the major source of new fronds every year, are thought not to undergo self-thinning during the growth season, unlike those from crowded stands of unitary seaweeds. For clonal seaweeds, it is not known, however, what happens at the very end of the growth season, when crowding is highest for the year. By sampling twice more frequently than previously done for similar species, the possible occurrence of frond self-thinning was tested for Mastocarpus papillatus (Rhodophyta, Gigartinales, Petrocelidaceae) from western Canada during the growth season (spring) of 2003. Initially, stand biomass increased together with frond density, as found previously for similar clonal seaweeds. Shortly before stand biomass peaked for the year (June), frond density remained statistically unchanged. Thus, the increased sampling precision of this study confirms that fronds of these clonal seaweeds do not undergo self-thinning, not even shortly before crowding is highest. Frond size inequality for M. papillatus remained statistically similar during the growth season, which is also consistent with a model of no self-thinning. There are similarities in biomass–density dynamics and in size inequality dynamics between clonal seaweeds and clonal vascular plants.     

Plant Interactions Alter the Predictions of Metabolic Scaling Theory

Metabolic scaling theory (MST) is an attempt to link physiological processes of individual organisms with macroecology. It predicts a power law relationship with an exponent of −4/3 between mean individual biomass and density during density-dependent mortality (self-thinning). Empirical tests have produced variable results, and the validity of MST is intensely debated. MST focuses on organisms’ internal physiological mechanisms but we hypothesize that ecological interactions can be more important in determining plant mass-density relationships induced by density. We employ an individual-based model of plant stand development that includes three elements: a model of individual plant growth based on MST, different modes of local competition (size-symmetric vs. -asymmetric), and different resource levels. Our model is consistent with the observed variation in the slopes of self-thinning trajectories. Slopes were significantly shallower than −4/3 if competition was size-symmetric. We conclude that when the size of survivors is influenced by strong ecological interactions, these can override predictions of MST, whereas when surviving plants are less affected by interactions, individual-level metabolic processes can scale up to the population level. MST, like thermodynamics or biomechanics, sets limits within which organisms can live and function, but there may be stronger limits determined by ecological interactions. In such cases MST will not be predictive.     

Effects of positive interactions, size symmetry of competition and abiotic stress on self-thinning in simulated plant populations

Background and Aims

Competition drives self-thinning (density-dependent mortality) in crowded plant populations. Facilitative interactions have been shown to affect many processes in plant populations and communities, but their effects on self-thinning trajectories have not been investigated.

Methods

Using an individual-based ‘zone-of-influence’ model, we studied the potential effects of the size symmetry of competition, abiotic stress and facilitation on self-thinning trajectories in plant monocultures. In the model, abiotic stress reduced the growth of all individuals and facilitation ameliorated the effects of stress on interacting individuals.

Key Results

Abiotic stress made the log biomass – log density relationship during self-thinning steeper, but this effect was reduced by positive interactions among individuals. Size-asymmetric competition also influenced the self-thinning slope.

Conclusions

Although competition drives self-thinning, its course can be affected by abiotic stress, facilitation and competitive symmetry.     

The effects of salt stress and arbuscular mycorrhiza on plant neighbour effects and self-thinning

Abiotic and biotic factors can alter the nature and strength of plant–plant interactions and therefore self-thinning (density-dependent mortality), but few studies have looked at how such factors interact. We investigated how salt stress and arbuscular mycorrhizal fungi (AMF) influence plant neighbour effects and self-thinning in experimental populations of Medicago sativa. We obtained two mycorrhizal levels by applying the fungicide benomyl (low AMF) or not (high AMF) at three salinity levels (0.05%, 0.2% and 0.5%). In experiment 1, we investigated how salinity and AMF interact to influence plant interaction intensity using a neighbour removal treatment. In experiment 2, we investigated how self-thinning dynamics vary under salinity conditions and different AMF levels at two initial plant densities (6000 and 17,500 seeds m^?2). Shoot biomass and plant density were measured 30, 60 and 90 days after sowing. Standardized major axis regression was used to estimate self-thinning parameters. In experiment 1, AMF increased competitive plant neighbour effects when there was no salinity stress, but this enhancement was not significant with increasing salinity. In experiment 2, there were effects of salinity and AMF on the self-thinning trajectory. The slope of the log (mean shoot biomass per unit area) vs. log density relationship was significantly steeper for the high AMF treatment than for the low AMF treatment without salinity, but the effect of AMF level on the self-thinning exponent was not significant under the two higher salinity levels. The effect of AMF treatments on the intercept of the self-thinning line was not significant at 0.2% salinity but was significant at 0.5% salinity, higher elevation for high AMF treatment. In self-thinning populations, AMF decreased the survival rate without salinity, but increased the survival rate at the highest salinity level. Our results support the hypothesis that salinity and AMF interact to influence plant neighbour effects and self-thinning. Under no-salinity conditions, AMF increased competition, steepened the self-thinning line and decreased survival rate, but these effects of AMF were not significant in the presence of salinity.     

Body size–density relationship for Mytilus edulis in an experimental food-regulated situation

We grew mussels ( Mytilus edulis ) under two different food regimes and eight population density levels to estimate the joint effects of density and biomass on their growth and survival and to determine the shape of the biomass–density ( B–N) relationship. Mussels were reared for 22 months, between December 1994 and October 1996, in 1-L experimental chambers supplied with natural seston. Growth in shell length, individual wet mass and ash free dry mass ( m ) decreased with decreasing food availability and increasing population density. Survival was negatively correlated with density but did not differ significantly between food regimes during the first year. Variations in concentration of available food did not alter the effects of crowding on mussels, as judged from the slopes of the body size–density curves. The general patterns exhibited by B–N curves were not consistent with expectations since we found 1) no classical competition–density (C–D) effect as reported in plants at intermediate competition levels, and 2) a slope of −0.648 for m–N curves in both food regimes, which was higher than expected from self-thinning (ST) theory. This value does not support present food-driven ST theory. This study introduces an unusual m–N region which combines properties of both ST and C–D effect.     

Effects of nutrient level on thinning and non-thinning crowding in even-aged populations of subterranean clover

Populations of subterranean clover were used to examine the effect of lowering nutrient supply on crowding in even-aged monospecific populations. Two hypotheses were being tested. Under one, termed ‘altered-speed’, reducing the nutrient supply to populations merely slows down the crowding process. Under the other, called ‘altered-form’, reducing the nutrient supply intensifies the crowding process (Morris & Myerscough 1984). The populations were grown at eight densities of sowing and three levels of nutrient supply. Non-thinning yield-density curves and self-thinning lines were fitted to the data. For the thinning populations, reducing the nutrient supply led to altered-form crowding, with thinning lines of reduced slope and intercept for total and shoot weight being observed in populations grown at lower levels of nutrient supply. For the non-thinning yield-density curves, lowering the nutrient supply did alter crowding effects, but not to the extent necessary to give full altered-form crowding. Rather, crowding effects intermediate between altered-speed and altered-form were observed. No altered-speed crowding was observed in this experiment. Comparison was made between the altered-speed crowding observed in other data (Hozumi & Ueno 1954: White & Harper 1970) with the altered-form crowding observed here and elsewhere (Hozumi & Ueno 1954). In these data, altered-form crowding was consistently associated with an increase in the proportion of root in the plants' growth as nutrient level was reduced, while in altered-speed crowding there was a complete lack of any such nutrient effect on root-shoot proportion. This was taken as evidence that under altered-speed crowding, variations in the level of nutrient supply did not lead to major change in the way that plants interfered with each other. For altered-form crowding, major change in the way plants interfered with each other's growth did occur as nutrient level fell.     

Competitive regulation of plant allometry and a generalized model for the plant self-thinning process

Taking into account the individual growth form (allometry) in a plant population and the effects of intraspecific competition on allometry under the population self-thinning condition, and adopting Ogawa's allometric equation 1/y = 1/axb + 1/c as the expression of complex allometry, the generalized model describing the change mode of r (the self-thinning exponential in the self-thinning equation, log M = K + log N, where M is mean plant mass, K is constant, and N is population density) was constructed. Meanwhile, with reference to the changing process of population density to survival curve type B, the exponential, r, was calculated using the software MATHEMATICA 4.0. The results of the numerical simulation show that (1) the value of the self-thinning exponential, r, is mainly determined by allometric parameters; it is most sensitive to change of b of the three allometric parameters, and a and c take second place; (2) the exponential, r, changes continuously from about -3 to the asymptote -1; the slope of -3/2 is a transient value in the population self-thinning process; (3) it is not a 'law' that the slope of the self-thinning trajectory equals or approaches -3/2, and the long-running dispute in ecological research over whether or not the exponential, r, equals -3/2 is meaningless. So future studies on the plant self-thinning process should focus on investigating how plant neighbor competition affects the phenotypic plasticity of plant individuals, what the relationship between the allometry mode and the self-thinning trajectory of plant population is and, in the light of evolution, how plants have adapted to competition pressure by plastic individual growth.     

Positive and Negative Interactions among Individuals of a Root Hemiparasite

Abstract: I studied the effects of a wide range of densities on establishment, survival, growth and reproduction of the annual root hemiparasite Rhinanthus alectorolophus in a field experiment. Seeds of the parasite were sown with those of a mixture of grassland plants as potential hosts. In most plants, seedling survival is strongly reduced by self-thinning at high densities, but in R. alectorolophus the proportion of seeds producing a young plant increased linearly with sowing density, indicating positive interactions among seedlings. Because survival to maturity was not influenced by density, the number of flowering plants per seed sown also increased with density. In contrast, mean plant size and reproduction were strongly reduced at high densities. It is suggested that resource sharing among parasites connected by haustoria is the most likely mechanism responsible for the reduced mortality of seedlings at high densities. The results indicate that facilitation among cohorts of conspecific root hemiparasites can increase the recruitment of young plants. The number of seeds produced per seed sown (a multiplicative fitness measure) was, however, independent of density in Rhinanthus because the early positive effects of density on recruitment were compensated later by the negative effects of crowding on growth and reproduction. Increased survival of seedlings could, however, indirectly increase fitness because it will increase the genetic diversity of offspring and may thus, for instance, reduce the impact of pathogens.     

Density effect, self-thinning and size distribution in Pinus densiflora Sieb. et Zucc. stands

The competition-density (C-D) effect for given times and self-thinning over time in even-aged, natural, pure stands of Pinus densiflora Sieb. et Zucc. were analyzed with the reciprocal equation of the C-D effect in self-thinning stands, and the equation describing the time-trajectory of mean stem volume and stand density. The C-D effect and self-thinning were consistently well explained by the two equations. Differences in mean stem volume and in stand density among the stands tended to merge with increasing stand age. The self-thinning line with a slope of approximately –3/2 was reached by the higher density stand prior to the medium and lower density stands. The skewness of tree height distribution showed positive values, which means that the distribution is more or less L-shaped, and in addition the skewness decreased with increasing mean tree height, which indicates that smaller trees died as the stands grew. This trend is consistent with the asymmetric (one-sided) competition hypothesis that self-thinning is driven by competition for light. The tree height distribution was analyzed using the Weibull distribution. The location parameter h _min of the Weibull distribution increased with increasing stand age, and the scale parameter a tended to increase slightly with increasing stand age. The range of the shape parameter b of the Weibull distribution corresponded to that of the skewness.     

Theoretical considerations on the C-D effect in self-thinning plant populations

A model for describing the competition–density (C-D) effect in self-thinning populations was developed on the basis of the following three basic assumptions: (1) the growth of mean phytomass follows a general logistic equation; (2) final yield is independent of initial population density; and (3) there exists a functional relationship between actual and initial population densities at any given time. The resultant equation takes the same reciprocal form as the reciprocal equation of the C-D effect derived from Shinozaki–Kira's theory (i.e., the logistic theory of the C-D effect), which deals with the density effect in nonself-thinning populations. However, one of the two time-dependent coefficients is quite different in mathematical interpretation between the two reciprocal equations. The reciprocal equation for self-thinning populations is essentially the same as the reciprocal equation assumed in the derivation of the functional relationship between actual and initial population densities. The establishment of the reciprocal equation is supported by the empirical facts that the reciprocal relationship between mean phytomass and population density is discernible in not only nonself-thinning populations but also in self-thinning populations. The present model is expected to systematically interpret underlying mechanisms between the C-D effect, which is observed at a time constant among populations with various initial densities, and self-thinning, which is observed along a time continuum in a given population. Received: August 5, 1998 / Accepted: January 7, 1999     

Density effects on organs in self-thinning <Emphasis Type="Italic">Pinus densiflora</Emphasis> Sieb. et Zucc. stands

Although much research on the density effect in nonself-thinning populations has been conducted, there has been very little research on density effects in self-thinning populations. Furthermore, the density effect of plant organs in self-thinning populations is little reported. The present study analyzed the yield–density (Y–D) effects on organs, such as stem, branch and leaf, together with that on stands of self-thinning Pinus densiflora Sieb. et Zucc.. The stand yield- and organ Y–D effects were well described by reciprocal and parabolic equations, respectively, throughout the experiment. The value of coefficient B in the reciprocal equation decreased monotonically with increasing stand age and became significantly closer to zero at the end of experiment (33-year-old stand), indicating that the constant final stand yield was established regardless of the density realized. The value of the relative growth coefficient h in the allometric equation between mean organ weight and mean aboveground weight was significantly smaller than 1.0 for stem, indicating that stem yield increases monotonically with increasing realized density. The h-value was significantly larger than 1.0 for branch throughout the experiment, and for leaf except at 33 years old, indicating that optimum densities exist. The h-value for leaf was not significantly different from 1.0 at 33 years old, indicating that the leaf yield reached a constant level regardless of realized density. The constant final leaf yield was established at almost the same growth stage as the establishment of constant final stand yield.     

Allometry of territory size and metabolic rate as predictors of self-thinning in young-of-the-year Atlantic salmon

1. Self-thinning is a progressive decline in population density caused by competitively induced losses in a cohort of growing individuals and can be depicted as: log₁₀ (density) = c − β log₁₀ (body mass).
2. In mobile animals, two mechanisms for self-thinning have been proposed: (i) the space hypothesis predicts that maximum population density for a given body size is the inverse of territory size, and hence, the self-thinning slope is the negative of the slope of the allometric territory-size relationship; (ii) the energetic equivalence hypothesis predicts that the self-thinning slope is the negative of the slope of the allometric metabolic rate relationship, assuming a constant supply of energy for the cohort.
3. Both hypotheses were tested by monitoring body size, population density, food availability and habitat for young-of-the-year Atlantic salmon ( Salmo salar ) in Catamaran Brook, New Brunswick. The results were consistent with the predictions of the space hypothesis. Observed densities did not exceed the maximum densities predicted and the observed self-thinning slope of −1·16 was not significantly different from the slope of −1·12, predicted by the allometry of territory size for the population under study.
4. The observed self-thinning slope was significantly steeper than −0·87, predicted by the allometry of metabolic rate, perhaps because of a gradual decline in food abundance over the study period. The decline in density was more rapid in very shallow sites and may have been partly caused by a seasonal change in water depth and an ontogenetic habitat shift rather than solely by competition for food or space.
5. The allometry of territory size may be a useful predictor of self-thinning in populations of mobile animals competing for food and space.     

Two‐phase self‐thinning in stream‐living juveniles of lake‐migratory brown trout Salmo trutta L. Compatibility between linear and non‐linear patterns across populations?

The self-thinning rule establishes that, in resource limited populations, increased per-capita requirements by individuals with indeterminate growth may limit density. This invariant rule relating log-log linear declines of cohort density with increased body mass has been well established in sessile organisms. However, few studies have assessed the occurrence of self-thinning in mobile organisms and still fewer have demonstrated its operation in natural situations. In Bisballe Baeck, where brown trout individuals experienced the simultaneous operation of density-dependence on growth and mortality, increased body mass explained 89.0–98.4% of the variations in density. However, relative to the invariant linear line, the decline in density with increased body mass described distinctly nonlinear, two-phase trajectories. An initial phase of variable slopes continued until body mass attained ≈10.2 g, at which the direction of the trajectory switched to a second phase of substantially steeper slopes or severe declines of density per mass unit increment. As inferred by the highly significant relationships between these two sets of slopes and annual recruitment, the two phases were consistent with a major assumption of self-thinning: the rate at which density declined with increased body mass increased with the intensity of intraspecific competition. The results of this study provide prima facie evidence for the occurrence of self-thinning in a mobile species based on observational data. A further comparison among density–body mass relationships described for resident, anadromous and lake-migratory brown trout suggests consistent two-phase self-thinning patterns across populations.     

Density effects on the growth of self-thinning Eucalyptus urophylla stands

Density effects on the growth of self-thinning Eucalyptus urophylla stands were examined for 7 years. Tree height and stem diameter at breast height were measured during the experimental period. Stems, branches, leaves, bark and roots of 45 E. urophylla trees were sampled in three different density stands in order to establish their biomass equations. Change trends of the biological time τ and density ρ were described used corresponding equations. The stem weight ratio increased and leaf weight ratio decreased, whereas those of branch, bark and root were relatively steady from 2 years after the planting. The competition-density (C-D) effect equation of mean organ weight w _o was derived by combining the allometric power relationship between mean tree weight w and w _o with the C-D effect equation of self-thinning stands. The equations of the C-D effect for w and ρ and for w _o and ρ were used to describe the C-D effects in tree and organs during course of self-thinning, respectively, and showed a good fit to the data. Leaf biomass of different density stands reached a more or less constant level with time elapse. High density produced the greatest biomass and stem biomass, so that it is the best choice in silvicultural practice.     

Arbuscular mycorrhizal fungi alter plant allometry and biomass-density relationships

Background and Aims

Plant biomass–density relationships during self-thinning are determined mainly by allometry. Both allometry and biomass–density relationship have been shown to vary with abiotic conditions, but the effects of biotic interactions have not been investigated. Arbuscular mycorrhizal fungi (AMF) can promote plant growth and affect plant form. Here experiments were carried out to test whether AMF affect plant allometry and the self-thinning trajectory.

Methods

Two experiments were conducted on Medicago sativa L., a leguminous species known to be highly dependent on mycorrhiza. Two mycorrhizal levels were obtained by applying benomyl (low AMF) or not (high AMF). Experiment 1 investigated the effects of AMF on plant growth in the absence of competition. Experiment 2 was a factorial design with two mycorrhizal levels and two plant densities (6000 and 17 500 seeds m⁻²). Shoot biomass, root biomass and canopy radius were measured 30, 60, 90 and 120 d after sowing. The allometric relationships among these aspects of size were estimated by standardized major axis regression on log-transformed data.

Key Results

Shoot biomass in the absence of competition was lower under low AMF treatment. In self-thinning populations, the slope of the log (mean shoot biomass) vs. log density relationship was significantly steeper for the high AMF treatment (slope = –1·480) than for the low AMF treatment (–1·133). The canopy radius–biomass allometric exponents were not significantly affected by AMF level, but the root–shoot allometric exponent was higher in the low AMF treatment. With a high level of AMF, the biomass–density exponent can be predicted from the above-ground allometric model of self-thinning, while this was not the case when AMF were reduced by fungicide.

Conclusions

AMF affected the importance of below-ground relative to above-ground interactions and changed root vs. shoot allocation. This changed allometric allocation of biomass and altered the self-thinning trajectory.     

How does fertility of the substrate affect intraspecific competition? Evidence and synthesis from self-thinning

When dense populations of even-aged plant monocultures are subject to intense competition, mortality can occur in a process known as self-thinning, in which changes in biomass are accompanied by decreases in density. On a plot of log biomass versus log density, self-thinning populations show a linear relationship called the self-thinning line. Variations in the fertility level of the substrate are known to affect self-thinning in a number of ways. Populations from substrates with different fertility levels have been observed to self-thin along the same line, or along different lines. A review of several experiments using the one species grown at different fertility levels was undertaken to look for any mechanisms that might account for the different patterns observed. It was postulated that the critical difference between whether populations followed a common or different line was the way in which competition developed in the stands as biomass accumulated. For the common-line pattern, data on the canopy volume required to support a given biomass showed that biomass packing did not differ between fertility levels, supporting the model of a common competitive mechanism operating at all fertility levels. When different lines were observed, the development of competition differed as plants increased in size and biomass accumulated at each fertility level. Over the upper range of fertility levels, biomass packing values per plant increased as fertility declined and the position of self-thinning lines followed predictions from biomass packing data. At the low end of the fertility scale, biomass packing values still decreased with fertility level, but the position of self-thinning lines was not linked to the biomass packing of individual plants: root interactions were presumed to dominate competition and the trajectory of self-thinning lines.     

RAMET DYNAMICS FOR THE CLONAL SEAWEED PTEROCLADIELLA CAPILLACEA (RHODOPHYTA): A COMPARISON WITH CHONDRUS CRISPUS AND WITH MAZZAELLA CORNUCOPIAE (GIGARTINALES)

Little is known about the dynamics and the ecological interactions among ramets (fronds) from populations of clonal red seaweeds. Small ramets are very difficult to tag, so their growth cannot be monitored directly. The temporal variation of the relationship between stand biomass and ramet density offers information on ramet performance. We calculated this relationship for an intertidal population of Pterocladiella capillacea (Gmelin) Santelices et Hommersand (Gelidiales) from Baja California, Mexico. Biomass and density were positively correlated on an annual basis, indicating that biomass accumulated without involving self-thinning among ramets. This contrasts with nonclonal seaweeds, for which self-thinning among individuals occurs during growth, but agrees with other clonal red seaweeds, such as Chondrus crispus Stackhouse and Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (both Gigartinales). The growth pattern for these members of the Gelidiales and of the Gigartinales holds despite differences in holdfast morphology and ramet branching degree and despite differences in the capacity of coalescence during early stages, known only for the Gigartinales. The positive slope for the dynamic biomass–density relationship, on a bilogarithmic scale, was statistically steeper for M. cornucopiae than for P. capillacea and for C. crispus. This suggests that the addition of new ramets during the growth season may be relatively more beneficial for biomass accumulation rates for M. cornucopiae. This would be expected for high-intertidal species subjected to strong abiotic stress, for which ramet crowding constitutes a key protection. Pterocladiella capillacea occurs at the mid-intertidal zone and C. crispus at the subtidal zone, so ramets would be relatively less important in that respect.     

Ecological and mathematical considerations on self-thinning in even-aged pure stands

A new model is proposed to unite the logistic theory of plant growth and the 3/2 power law of self-thinning, which so far have been applied independently to growth analysis. To construct the model the following assumptions are made: a general logistic curve of mean plant weight, a modified form of the formula to show the rule of constancy of the final yield, which is generalized to cover the conditions of different combinantions of density and linear factor supply in a nonself-thinning population and a special population with a specific initial density which follows thew-ρ trajectory of the 3/2 power law type and has an exponential decrease in its density with biological time. Model calculations show that the Sukatschew effect is successfully formulated, that there should be a minimum factor supply below which self-thinning does not occur and that thew-ρ trajectory should be segregated acoording to the level of the linear factor supply.     

Ecological and mathematical considerations on self-thinning in even-aged pure stands

It is emphasized in growth analysis of self-thinning populations that relative mortality rate pertains to the difference between relative growth rates and net assimilation rates, each of which are definable on a mean plant size basis or on a biomass basis. The time trends of the ratio of relative mortality rate to relative growth rates to be expected according to Tadaki's, Shinozaki's and Hozumi's models are compared with that of the eastern white pine population, and a good agreement is exhibited. As an alternative to Hozumi's model, a new model is constructed to unite the logistic theory of plant growth and the 3/2 power law concerning self-thinning, which so far have usually been applied independently to growth analysis. To construct the model the following assumptions are made: the fundamental equation to relate mean plant weight with density in self-thinning population proposed by Shinozaki, and a special population with a specific initial density which follows thew-p trajectory of the 3/2 power law type and has an exponential decrease in its density with biological time. Properties of the model are examined from ecological and mathematical viewpoints.     

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.