Plants in a crowded stand regulate their height growth so as to maintain similar heights to neighbours even when they have potential advantages in height growth

Background and Aims

Although being tall is advantageous in light competition, plant height growth is often similar among dominant plants in crowded stands (height convergence). Previous theoretical studies have suggested that plants should not overtop neighbours because greater allocation to supporting tissues is necessary in taller plants, which in turn lowers leaf mass fraction and thus carbon gain. However, this model assumes that a competitor has the same potential of height growth as their neighbours, which does not necessarily account for the fact that height convergence occurs even among individuals with various biomass.

Methods

Stands of individually potted plants of Chenopodium album were established, where target plants were lifted to overtop neighbours or lowered to be overtopped. Lifted plants were expected to keep overtopping because they intercept more light without increased allocation to stems, or to regulate their height to similar levels of neighbours, saving biomass allocation to the supporting organ. Lowered plants were expected to be suppressed due to the low light availability or to increase height growth so as to have similar height to the neighbours.

Key Results

Lifted plants reduced height growth in spite of the fact that they received higher irradiance than others. Lowered plants, on the other hand, increased the rate of stem elongation despite the reduced irradiance. Consequently, lifted and lowered plants converged to the same height. In contrast to the expectation, lifted plants did not increase allocation to leaf mass despite the decreased stem length. Rather, they allocated more biomass to roots, which might contribute to improvement of mechanical stability or water status. It is suggested that decreased leaf mass fraction is not the sole cost of overtopping neighbours. Wind blowing, which may enhance transpiration and drag force, might constrain growth of overtopping plants.

Conclusions

The results show that plants in crowded stands regulate their height growth to maintain similar height to neighbours even when they have potential advantages in height growth. This might contribute to avoidance of stresses caused by wind blowing.     

Density Effects on Plant Height Growth and Inequality in Sunflower Populations

Comparisons between competing and non-competing sunflower （Helianthus annuus L.） populations demonstrate pronounced effects of density on plant height growth, height-to-crown width ratio, and s popuiaUon＇s height inequality. In the present study, non-destructive measurements of height and the prolected crown area of sunflower plants were taken at seven times from emergence to fruit maturation in even-aged monospeclflc stands with initial densities of 1, 4, 16, and 64 plants/m^2. The mean height of populations Increased and then decreased with increasing population density; the height Inequalities of uncrowded populations decreased during stand growth, whereas the height inequaiiUes of crowded popuisUons decreased first and then increased during stand development. The interindlvidual relationships between the relative height growth rate and height within uncrowded populations became significantly negative during population growth, whereas these relationships were negative first and then became positive during the development of crowded populations. In the uncrowded populations, the static Interindlvldual relationship between height-to-crown width ratio and volume was positive, whereas for the crowded population these relationships became negative with increasing competition for light. The data suggest that the plastic responses of plant height and height-to-crown width ratio to light competition will become more Intense with increasing competition Intensity. The results of the present study argue strongly for the Importance of size-dependent Individual-level plastic responses due to size-asymmetric light competition In generating the variations in population height inequality.     

Using virtual plants to analyse the light-foraging efficiency of a low-density cotton crop

BACKGROUND AND AIMS: Cotton shows a marked plasticity vs. density in terms of branch development and geometry, internode elongation and leaf expansion. This paper proposes interpretations for observed plasticity in terms of light quantity and quality. METHODS: 3-D virtual plants were reconstructed from field observations and 3-D digitization and were used to simulate the light regime in cotton stands of different densities. KEY RESULTS: All densities showed the same linear relationship between LAI and the sum of light intercepted by the canopy, from seedling emergence up to flowering. Simulated R : FR ratio profiles can very likely explain (1) the longer first internodes on main stem and branches and (2) the azimuthal re-orientation of branches toward the inter-row. CONCLUSIONS: Simulation tools were used to analyse plant plasticity in terms of light quantity and quality. The methodology applied here at the stand scale will now be continued at the plant scale to further strengthen the above hypotheses.     

Height convergence in response to neighbour growth: genotypic differences in the stoloniferous plant Potentilla reptans

Using a new experimental set up, the way in which height growth of stoloniferous plants is adjusted to that of their neighbours, as well as differences between genotypes in their ability to keep up with neighbour height growth were tested. Five Potentilla reptans genotypes inherently differing in petiole length were subjected to three experimental light gradients, involving light intensity and red : far-red ratio. Each plant was placed in a vertically adjustable cylinder of green foil, and the treatments differed in the speed of cylinder height increase and final height. Total weight of plants decreased from the 'Slow' to the 'Fast' treatment, while petiole length increased. Leaves reaching the top of the cylinder stopped petiole elongation, resulting in similar final heights for all genotypes in the 'Slow' treatment. In the 'Fast' treatment only the fastest-growing genotype maintained its position in the top of the cylinder and genotypes differed strongly in final height within the cylinders. Plants adjust their height growth to that of the surrounding vegetation, leading to height convergence in short light gradients that slowly increase. These adjustments and genotypic differences in ability to keep up with fast-growing neighbours can influence the outcome of competition for light.     

Photocontrol of stem elongation in plant neighbourhoods: effects of photon fluence rate under natural conditions of radiation

Abstract. We investigated the effects of photon fluence rate on internode elongation in fully de-etiolated plants growing under sunlight. Our goal was to find out whether perception by the stems of fluence rate changes related to canopy density may be involved in the modulation of internode growth in canopies formed by plants of similar stature (e.g. crop stands). Using Datura ferox L. and Sinapis alba L. seedlings growing under natural radiation, we found that internode elongation is promoted by localized shading. This effect was observed with internodes receiving light with a high (>0.9) or a low (0.3) red (R) to far-red (FR) photon ratio. Selective removal of the different wavebands from the light impinging on the internodes showed that part of the response to fluence rate is due to photons in the R + FR range. The blue (B) component, most likely acting through a specific photoreceptor, also inhibited elongation. However, changes in the fluence rate of B light did not have detectable effects on the response of the internodes to R:FR ratio. Fibre-optic studies and measurements with integrating-cylinder sensors in even-aged populations of seedlings showed that both the quality and quantity of radiation received by the stems are profoundly influenced by changes in canopy density. When density is very low (leaf area index = LAI ≥ 1) only the R:FR ratio is reduced, due to FR reflected from nearby leaves. In the LAI range of 1 to 2, though a large proportion of the leaf area is still receiving full sunlight, the photon fluence rate at the stem level drops dramatically. These results suggest that in even-aged populations of LAI > 1 elongation growth is promoted by the low R:FR ratio and the reduced fluence rate. Perception of these two factors at the stem level may elicit morphological adaptations in the canopy before the onset of severe competition among neighbours for the resource of light.     

Density-induced plant size reduction and size inequalities in ethylene-sensing and ethylene-insensitive tobacco

Abstract: Plant competition for light is a commonly occurring phenomenon in natural and agricultural vegetations. It is typically size-asymmetric, meaning that slightly larger individuals receive a disproportionate share of the light, leaving a limited amount of light for the initially smaller individuals. As a result, size inequalities of such stands increase with competition intensity. A plant's ability to respond morphologically to the presence of neighbour plants with enhanced shoot elongation, the so-called shade avoidance response, acts against the development of size inequalities. This has been shown experimentally with transgenic plants that cannot sense neighbours and, therefore, show no shade avoidance responses. Stands of such transgenic plants showed a much stronger development of size inequalities at high plant densities than did wild type (WT) stands. However, the transgenic plants used in these experiments displayed severely hampered growth rates and virtually no response to neighbours. In order to more precisely study the impact of this phenotypic plasticity on size inequality development, experiments required plants that have normal growth rates and reduced, but not absent, shade avoidance responses. We made use of an ethylene-insensitive, transgenic tobacco genotype (Tetr) that has wild type growth rates and moderately reduced shade avoidance responses to neighbours. Here, we show that the development of size inequalities in monocultures of these plants is not affected unambiguously different from wild type monocultures. Plots of Tetr plants developed higher inequalities for stem length than did WT, but monocultures of the two genotypes had identical CV (Coefficient of Variance) values for shoot biomass that increased with plant density. Therefore, even though reduced shade avoidance capacities led to the expected higher size inequalities for stem length, this does not necessarily lead to increased size inequalities for shoot biomass.     

The effects of cassava mosaic virus disease on yield and compensation in mixed stands of healthy and infected cassava

The effects of cassava mosaic virus disease (CMD) on yield in fully and partly infected stands of cassava were investigated in field trials in Uganda in 1990-91 and 1991-92. Three cultivars (Ebwanateraka, Bao and Bukalasa 1 l), each at three levels of cutting infection (O%, 50% and 100%) and harvested 510 and 15 months after planting (MAP) were used in a randomised block design with split-split plots and four replicates. Moreover, yield and growth data for individual infected and uninfected plants were considered in relation to the health status of their nearest neighbours. In each experiment, fresh tuberous root yields of plants from 100% infected plots gave sigdicantly lower yields than those from 0% or 50% infected plots at each harvest date and the losses were greatest in cv. Bao. Yields of plants from 0% and 50% plots for each of the three cultivars were not significantly different, 10 and 15 MAP. The loss in yield differed between cultivars and harvest dates. Fresh stem, leaf and root yields and the number of tuberous roots were influenced by the health status of the plants harvested and that of their nearest neighbours. Uninfected plants surrounded by infected ones had more roots and heavier total fresh root, stem and leaf weights than those surrounded by uninfected ones. Overall, 26% and 42% compensation was recorded in 1990-91 and 1991-92, respectively. The effects of CMD on cassava production and of compensation in mixed stands of infected and uninfected plants are discussed, especially in relation to control strategies such as roguing.     

Ethylene is required in tobacco to successfully compete with proximate neighbours

Plants sense neighbours even before these cause a decrease in photosynthetic light availability. Light reflected by proximate neighbours signals a plant to adjust growth and development, in order to avoid suppression by neighbour plants. These phenotypic changes are known as the shade‐avoidance syndrome and include enhanced shoot elongation and more upright‐positioned leaves. In the present study it was shown that these shade‐avoidance traits in tobacco (Nicotiana tabacum) are also induced by low concentrations of ethylene. Furthermore, it was shown that transgenic plants, insensitive to ethylene, have a delayed appearance of shade‐avoidance traits. The increase in both leaf angles and stem elongation in response to neighbours are delayed in ethylene‐insensitive plants. These data show that ethylene is an important component in the regulation of neighbour‐induced, shade‐avoidance responses. Consequently, ethylene‐insensitive plants lose competition with wild‐type neighbours, demonstrating that sensing of ethylene is required for a plant to successfully compete for light.     

Tree shape plasticity in relation to crown exposure

Trees outside closed forest stands differ in the relation between stem diameter, height and crown volume from trees that grew with neighbours close by. Whether this plasticity in tree shape varies between species in relation to their light requirement is unknown. We purposefully sampled 528 trees ranging 5–100?cm diameter at breast height growing in a range of light conditions. Across ten broad-leaved species observed in Sumatra or Kalimantan, a generic relationship was found between light exposure of the crown and a light-dependent a _l parameter that modifies the height–diameter allometric equation (H?=?a _l D ^b ) from those for closed stands. In our results, vertical stretching is well predicted by light availability. In fully open conditions, trees are on average 31% shorter for the same diameter than under (partial) shade. Most of the stretching response occurs in all species as soon as some degree of lateral shading occurs. The response, however, varies by species (8–44% reduction) in a way apparently unrelated to species’ successional status. Crown volume varied less than stem height in its relationship with stem diameter across all light conditions tested. The scaling of crown volume with stem diameter, however, differed markedly between tree species.     

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     

Influence of elevated CO2 on canopy development and red:far-red ratios in two-storied stands ofRicinus communis

Vertical structure of plant stands and canopies may change under conditions of elevated CO₂ due to differential responses of overstory and understory plants or plant parts. In the long term, seedling recruitment, competition, and thus population or community structure may be affected. Aside from the possible differential direct effects of elevated CO₂ on photosynthesis and growth, both the quantity and quality of the light below the overstory canopy could be indirectly affected by CO₂-induced changes in overstory leaf area index (LAI) and/or changes in overstory leaf quality. In order to explore such possible interactions, we compared canopy leaf area development, canopy light extinction and the quality of light beneath overstory leaves of two-storied monospecific stands ofRicinus communis exposed to ambient (340 μl l⁻¹) and elevated (610 μl l⁻¹) CO₂. Plants in each stand were grown in a common soil as closed “artificial ecosystems” with a ground area of 6.7 m². LAI of overstory plants in all ecosystems more than doubled during the experiment but was not different between CO₂ treatments at the end. As a consequence, extinction of photosynthetically active radiation (PAR) was also not altered. However, under elevated CO₂ the red to far-red ratio (R:FR) measured beneath overstory leaves was 10% lower than in ecosystems treated with ambient CO₂. This reduction was associated with increased thickness of palisade layers of overstory leaves and appears to be a plausible explanation for the specific enhancement of stem elongation of understory plants (without a corresponding biomass response) under elevated CO₂. CO₂ enrichment led to increased biomass of overstory plants (mainly stem biomass) but had no effect on understory biomass. The results of this study raise the possibility of an important indirect effect of elevated CO₂ at the stand-level. We suggest that, under elevated CO₂, reductions in the R:FR ratio beneath overstory canopies may affect understory plant development independently of the effects of PAR extinction.     

Effects of mechanical stress and plant density on mechanical characteristics, growth, and lifetime reproduction of tobacco plants

Plastic increases in stem elongation in dense vegetation are generally believed to be induced by canopy shading, but because plants protect each other from wind, shielding (reduced mechanical stress) could also play a role. To address this issue, tobacco Nicotiana tabacum plants were subjected to two levels of mechanical stress, 0 (control) or 40 (flexed) daily flexures, and grown solitarily, in a dense monostand (with plants of only one mechanical treatment), or in a mixed stand (flexed and control plants grown together). Flexed plants produced shorter and thicker stems with a lower Young's modulus than control plants, while dense-stand plants had relatively taller and thinner stems than solitary ones. Flexing effects on stem characteristics were independent of stand density. Growth, reproduction, and survival of solitary plants were not affected by flexing, while in the monostand growth was slightly reduced. But in the mixed stand, flexed plants were readily shaded by controls and had considerably lower growth, survival, and reproduction rates. These results suggest that wind shielding indeed plays a role in the plastic increase in stem elongation of plants in dense vegetation and that this response can have important consequences for competitive ability and lifetime seed production.     

Effect of Canopy Structure on Degree of Asymmetry of Competition in Two Forest Stands in Northern Japan

The canopy structure in terms of the vertical distribution of leaf mass and the degree of asymmetry of competition between individual trees was studied in two types of forest stand in Hokkaido, northern Japan: a naturally regenerated stand ofBetulaspp. and an artificial plantation ofPicea abies.The canopy structure in theBetulastand was more hierarchical; larger individuals were not heavily shaded even in the lowest part of their crowns and smaller individuals were heavily shaded by their larger neighbours. The canopy structure in thePiceastand was less hierarchical; even larger individuals were shaded in the lowest part of their crowns and smaller individuals were not heavily shaded by their neighbours. Application of the general formula of size-dependent mean growth rate revealed that competition in theBetulastand was more one-sided than that in thePiceastand. This result was consistent with the trends in the change over time in size equality in both stands.Even if competition is mediated by light, which often makes competition one-sided, the degree of one-sidedness in competition can be variable depending on canopy structure.     

Stem Elongation Response to Neighbour Shade in Sprawling and Upright Polygonum Species

Shading by neighbouring plants, which reduces energy for photosynthesisand lowers the ratio of red:far red light, can trigger a stemelongation or ‘overtopping’ response in herbaceousplants. We compared the stem elongation response of two Polygonumspecies in a greenhouse experiment. P. sagittatum, a sprawling,vine-like annual, and P. hydropiperoides, an upright perennial,were grown from seeds at three levels of neighbour shade producedby crowding a cohort of real neighbour plants or adult-sizedfake neighbour plants that provided shade and reduced the red:far-redratio. We hypothesized that the annual would show a more pronouncedelongation response to developing or adult neighbour shade becausevine-like plants are less mechanically constrained to remainupright and self-supporting. Internodes on stems of bothP. sagittatumand P. hydropiperoides increased in length as the amount ofshading by real or fake neighbours increased. P. sagittatumclimbed on adjacent plants, and had longer stems with more nodesthan those of P. hydropiperoides. Although both P. sagittatumand P. hydropiperoides tended to elongate with crowding, thegreater elongation response to both real and fake neighboursshown by the sprawling annual reflects its ability to extendupward into a canopy beyond self-supporting height. Self-supportingP. hydropiperoides can extend upward with, or overtop, cohortneighbours, but might less readily elongate into an extant canopy.In dense stands, P. sagittatum can become structurally dependenton close P. hydropiperoides neighbours. Our results suggestthat the elongation response of P. sagittatum to neighbourscan contribute to structural dependence and could facilitatecoexistence of these species.Copyright 2000 Annals of BotanyCompany Neighbour effects, light variation, red:far-red, plant growth strategy, Polygonum sagittatum, Polygonum hydropiperoides, smartweeds     

Maintenance cost, toppling risk and size of trees in a self-thinning stand

Wind routinely topples trees during storms, and the likelihood that a tree is toppled depends critically on its allometry. Yet none of the existing theories to explain tree allometry consider wind drag on tree canopies. Since leaf area index in crowded, self-thinning stands is independent of stand density, the drag force per unit land can also be assumed to be independent of stand density, with only canopy height influencing the total toppling moment. Tree stem dimensions and the self-thinning biomass can then be computed by further assuming that the risk of toppling over and stem maintenance per unit land area are independent of stand density, and that stem maintenance cost is a linear function of stem surface area and sapwood volume. These assumptions provide a novel way to understand tree allometry and lead to a self-thinning line relating tree biomass and stand density with a power between −3/2 and −2/3 depending on the ratio of maintenance of sapwood and stem surface.     

Canopy studies on ethylene-insensitive tobacco identify ethylene as a novel element in blue light and plant-plant signalling

Plants growing at high densities express shade avoidance traits as a response to the presence of neighbours. Enhanced shoot elongation is one of the best researched shade avoidance components and increases light capture in dense stands. We show here that also leaf movements, leading to a more vertical leaf orientation (hyponasty), may be crucial in the early phase of competition. The initiation of shade avoidance responses is classically attributed to the action of phytochrome photoreceptors that sense red:far-red (R:FR) ratios in light reflected by neighbours, but also other signals may be involved. It was recently shown that ethylene-insensitive, transgenic (Tetr) tobacco plants, which are insensitive to the gaseous plant hormone ethylene, have reduced shade avoidance responses to neighbours. Here, we report that this is not related to a reduced response to low R:FR ratio, but that Tetr tobacco plants are unresponsive to a reduced photon fluence rate of blue light, which normally suppresses growth inhibition in wild-type (WT) plants. In addition to these light signals, ethylene levels in the canopy atmosphere increased to concentrations that could induce shade avoidance responses in WT plants. Together, these data show that neighbour detection signals other than the R:FR ratio are more important than previously anticipated and argue for a particularly important role for ethylene in determining plant responses to neighbours.     

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     

Biomechanical Plasticity Facilitates Invasion of Maritime Forests in the southern USA by Brazilian pepper (Schinus terebinthifolius)

Biomechanical plasticity and within-species growth form diversity are traits that can facilitate invasion by non-native plant species. We support this argument with evidence from the invasion of coastal habitats in northern Florida, USA, by Schinus terebinthifolius and describe some of the consequences of this invasion for overtopped saltmarsh plants. In crowded stands, Schinus grows more like a vine than a tree, with stem height : diameter ratios nearly twice than those observed in open-grown individuals but with no changes in wood density or the modulus of elasticity of stem material. When extracted from the surrounding vegetation, the formerly crowded Schinus stems buckle under their own weight. Schinus crowns also extend much further over adjacent saltmarsh than crowns of Juniperus virginiana, the only other tree species abundant in the study site. Along forest edges, the above-ground biomass of saltmarsh plants overtopped by Schinus crowns was reduced by more than an order of magnitude. The biomechanical plasticity of Schinus allows it to adapt its growth form to suit habitat conditions and can dominate the edges of salt marshes as a sprawling shrub and maritime forests as either a free-standing tree or a woody vine, depending on stand crowding     

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.     

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

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