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.     

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.     

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.     

Density-dependent Mortality Induced by Low Nutrient Status of the Substrate

The hypothesis that changing the fertility level of the substratewould change the self-thinning line (different slope or intercept)followed by high-density populations was tested by sowing populationsof Ocimum basilicum L. at two densities on a soil-based pottingmix adjusted to three fertility levels (F0, F1 and F2). Fertilitylevel significantly affected the slope of the thinning linesfor both shoot and root biomass. For shoot biomass, more mortalityoccurred per unit increase in biomass as fertility level declined(the slope of the thinning line became flatter). The slope ofthe log shoot biomassvs. log density relationship was -0.5 atthe F2-, zero at the F1-, and 0.94 at the F0-fertility. Forthe log root biomassvs. log density lines, slopes were zeroat the F2- and F0-fertility levels, and -0.32 at F1. Packingof shoot biomass into canopies of individual plants correlatedwell with observed exponents of self-thinning lines at the F2-and F1-fertility level. Plants at the F2-fertility level requiredmore canopy space to support a given shoot biomass than plantsat F1, indicating that shoot competition was more intense atthe F2-fertility level for a given biomass. Leaf area indexand size inequality also increased with fertility level fora given shoot biomass. Density-dependent mortality in populationsgrown at the F0-fertility level was highly unusual in havinga positive slope for the shoot biomass vs. density relationship.Shoot growth per plant was static as density declined in theF0-populations; however, root growth per plant increased. Allmeasurements of shoot growth (mass, height, canopy extension,leaf area) remained static in the F0-populations: root massand length increased in comparison. It is argued that root competitionbecame sufficiently intense to cause the density-dependent mortalityseen at the F0-fertility level, with little contribution ofshoot competition to mortality. Copyright 1999 Annals of BotanyCompany Ocimum basilicum, self-thinning, root competition, shoot competition, fertility level and competition, density-dependent mortality, allometric self-thinning.     

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.     

植物种群自疏过程中构件生物量与密度的关系

不论是在对植物种群自疏规律还是在对能量守衡法则的研究中,个体大小(M)大多针对植物地上部分生物量,地下部分和构件生物量及其动态十分重要又多被忽视。以1年生植物荞麦为材料研究了自疏种群地下部分生物量、包括地下部分的个体总生物量以及各构件生物量与密度的关系。结果表明:平均地上生物量和个体总生物量与密度的异速关系指数(γabove-ground和γindividual)分别为-1.293和-1.253,与-4/3无显著性差异(P>0.05),为-4/3自疏法则提供了有力证据;平均根生物量-密度异速指数γroot(-1.128)与-1无显著性差异(P>0.05),与最终产量恒定法则一致;平均茎生物量-密度异速指数γstem(-1.263)接近-4/3(P>0.05),平均叶生物量-密度异速指数γleaf(-1.524)接近-3/2(P>0.05),分别符合-4/3自疏法则与-3/2自疏法则;而繁殖生物量与密度的异速关系指数γreproductive(-2.005)显著小于-3/2、-4/3或-1(P<0.001)。因此,不存在一个对植物不同构件普适的生物量-密度之间的关系。光合产物在地上和地下构件的生物量分配格局以及构件生物量与地上生物量之间特异的异速生长关系导致不同构件具有不同的自疏指数。无论对于地上生物量还是个体总生物量,荞麦种群能量均守衡,而对于地下生物量,荞麦种群能量不守衡。     

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.     

-3/2方自疏法则的机理与普适性

根据植物构件性理论,对经验规律-3/2方自疏法则进行了理论推导,对-3/2方自疏法则以前不能解释的实验结果进行了理论分析,并以春小麦（Triticum aestiunm L.）为材料,设置了4个密度梯度的实验,定期测定个体形态及种群数量指标,证明了-3/2方自疏法法则对春小麦作物种群的适用性,同时,基于所测定的植株形态数据,结合数值的量纲变化特点对-3/2方自疏法则的普适性进行了理论解释。     

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.     

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     

Allocation, plasticity and allometry in plants

Allocation is one of the central concepts in modern ecology, providing the basis for different strategies. Allocation in plants has been conceptualized as a proportional or ratio-driven process (‘partitioning’). In this view, a plant has a given amount of resources at any point in time and it allocates these resources to different structures. But many plant ecological processes are better understood in terms of growth and size than in terms of time. In an allometric perspective, allocation is seen as a size-dependent process: allometry is the quantitative relationship between growth and allocation. Therefore most questions of allocation should be posed allometrically, not as ratios or proportions. Plants evolve allometric patterns in response to numerous selection pressures and constraints, and these patterns explain many behaviours of plant populations.

In the allometric view, plasticity in allocation can be understood as a change in a plant's allometric trajectory in response to the environment. Some allocation patterns show relatively fixed allometric trajectories, varying in different environments primarily in the speed at which the trajectory is travelled, whereas other allocation patterns show great flexibility in their behaviour at a given size. Because plant growth is often indeterminate and its rate highly influenced by environmental conditions, ‘plasticity in size’ is not a meaningful concept. We need a new way to classify, describe and analyze plant allocation and plasticity because the concepts ‘trait’ and ‘plasticity’ are too broad. Three degrees of plasticity can be distinguished: (1) allometric growth (‘apparent plasticity’), (2) modular proliferation and local physiological adaptation, and (3) integrated plastic responses. Plasticity, which has evolved because it increases individual fitness, can be a disadvantage in plant production systems, where we want to optimize population, not individual, performance.     

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.     

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.     

Plant Growth Analysis: Allometry, Growth and Interference in Orchardgrass and Timothy

A multiple regression procedure was used to evaluate allometricresponses to stand age and species population densities in monoculturesand mixtures of orchardgrass (Dactylis glomerata L., also knownas cocksfoot) and timothy (Phleum pratense L.). In each speciesthe allometry between shoot dry weight and either leaf areaor tiller number per plant was studied. Population density treatmentsaffected allometry by changing allometric exponents expressingthe ratio of relative growth rates of different plant characteristics.Allometric relationships changed as growth proceeded, and thetwo species differed in their allometric responses to treatments. Plant growth analysis, allometry, competition, Dactylis glomerata L., Phleum pratense L.     

植物的表型可塑性、异速生长及其入侵能力

表型可塑性是指同一个基因型对不同环境响应产生不同表型的特性,特定性状的可塑性本身可以遗传,也可以接受选择而发生进化。植物个体的异速生长是指生物体某一特征的相对生长速率不等于第二种特征的相对生长速率的特性,该特性是由物种的遗传性决定的一种固定特征,植物往往朝着最佳的异速生长曲线进化。植物特定基因型在不同环境下,诸如生物量分配和种群几何学上的一些表型差异,既可由异速生长造成,也可由表型可塑性造成。植物本身的异速生长是一种"外观可塑性",而异速生长曲线的改变才是真正的可塑性。植物的表型可塑性、异速生长对于入侵植物的适应具有重要意义。干扰等异质性生境下表型可塑性成为物种生存扩散的有利性状,表型可塑性强的物种更有可能成为广布种。植物本身的异速生长特性或其异速生长曲线的改变都能影响其入侵能力。     

Density-dependence in single-species populations of plants

The general form of yield-density relationships in plant populations is discussed with reference to reciprocal equations and the $\text{3}\text{2}$ power law, which describes the concomitant changes in plant weight and density during self-thinning. A model to describe the pattern of mortality in high density populations is also discussed with particular reference to the nature of intraspecific competition within plant populations.A reparameterized version of a reciprocal equation proposed by Bleasdale & Nelder is used to describe the relationship between individual plant weight and surviving plant density. The biological interpretation of the parameters is discussed in relation to the dry matter production of isolated plants, the density at which mutual interference between neighbours becomes appreciable, and the efficiency of resource utilization at high densities.The reparameterized equation is then used together with an equation which describes mortality during self-thinning as the basis for a new model to describe the relation between total plant yield and sowing density. The law of allometry is used in conjunction with the model to describe the relationship between the weight of a plant part and density, and this then forms the basis for a model of the population dynamics of annual plants with effectively discrete generations. Finally the dynamical behaviour of plant populations is discussed. It is concluded that most plant populations will show neighbourhood stability with exponential or perhaps oscillatory damping towards an equilibrium.     

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.     

The self-thinning process in mangrove <Emphasis Type="Italic">Kandelia obovata</Emphasis> stands

The self-thinning process was monitored in crowded Kandelia obovata Sheue, Liu & Yong stands over four years. The frequency distribution of tree phytomass was an L-shape, which was kept over the experimental period. Spearman’s rank correlation coefficient for phytomass decreased as the time span of the comparison became longer, a result which indicates that the rank of phytomass changes as stands grow. Death of trees resulted from one-sided competition, i.e., death occurred in lower-rank trees. Surviving trees continued to grow. Whatever the current spatial distribution of the trees, death occurred randomly and the spatial distribution gradually became close to random as stands grew. The self-thinning exponent was 1.46, which can be regarded as evidence in favor of the 3/2 power law of self-thinning. Relative growth rate, RGR, decreased in proportion to decreasing relative mortality rate, RMR, with a proportionality constant of 1.57, which was not significantly different from the slope of the self-thinning exponent. This experimental result probably justifies the assumption that the ratio of RGR to RMR in the mean phytomass-density trajectory for any self-thinning population with different densities becomes constant as the growth stage progresses.     

Sprout recruitment and self-thinning of Erica multiflora after clipping

Regrowth after clipping and the effect of local competition were studied in a natural population of Erica multiflora in a Mediterranean shrubland, by removing neighbours at 1 and 2 m around the target plants during four growing seasons. Removal of surrounding natural vegetation increased the number, the density (number of sprouts per stump area) and the biomass of the sprouts growing from clipped plants. Target plants ònly interacted with their near neighbours. Target plants had a negative relative increment in the number of sprouts per stump during the 18 months immediately following treatment, but a positive increment thereafter, which suggests that there was a constant or episodic recruitment of sprouts within the stump after clipping. Competition treatment had a non-significant effect on the negative increment of sprouts per stump. The self-thinning trajectory was different for the different competition treatments: there was an allometric negative relationship between density of sprouts and mean biomass of survivors during all sampling periods in genets without neighbours in a 1-m radius; the self-thinning trajectory of sprouts in genets without neighbours in a 2-m radius was short, a net increase in sprouts per stump area was accompanied by an increase in mean sprout biomass 30 months after clipping. During the same period, however, plants with neighbours showed a decline in both the sprout biomass and density.     

Plant self-thinning dynamics

Plant self-thinning of Hellianthus annuus is examined and it is shown that the mean leaf area ratio is equal to the mean plant density ratio to the power-4/3 independent of the mean plant dry weight and independent of the light intensity over the experimental range considered. The constant term of this basic self-thinning equation is identified, in terms of the mean leaf area and mean plant density values (L _c and p _c respectively) for the plant population in its earliest competing post germination stage and in terms of the derivatives (d log L/d log p)₁=(d log L/dt)₁/(d log p/dt)₁=-4/3 which is independent of light intensity, as L _c p_c ^-4/3.