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.     

Growth and competition in clonal plants-persistence of shoot populations and species diversity

This paper reviews studies on growth and size-structure dynamics of shoots and clones in clonal plants in comparison with those in non-clonal plants, and discusses the characteristics of clonal plants. The mode of competition between individuals (symmetric versus asymmetric, degree of competitive asymmetry), growth dynamics of individuals, allocation pattern between organs and spatial pattern of individuals are closely correlated with each other in non-clonal plant populations. Theoretical and field studies based on the diffusion model revealed that plants of “height-growth” type (mostly early-successional tree species) and plants of “diameter-growth” type (mostly late-successional tree species) tend to exhibit asymmetric competition and symmetric competition respectively. Moreover, asymmetrically competing plants show smaller effects of variation in individual growth rate and spatial pattern on the size-structure dynamics of the population than symmetrically competing plants. Thefefore, the spatial pattern of inviduals should be considered especially for plants undergoing symmetric competition. These results for non-clonal plants should have a significant implication also for the growth dynamics and competition in clonal plants. The mean growth rate of shoots [G(t,x) function] and hence the mode of competition between shoots differs among clonal plant species as in non-clonal plants. However, a large magnitude and size-independence (or slightly negative size-dependence) of the variation in growth rate of shoots [D(t,x) function], especially at the early stage in a growing season is a common characteristic of many clonal plant species, in contrast to the positively size-dependent variation in individual growth rate in non-clonal plants. This type of variation in shoot growth rate leads to the persistence of stable shoot populations even when the mean growth rate function is changed, and also in cases where the shoot population structure would be unstable in the absence of variation in growth rate. It is suggested that competition between clones is symmetric in most clonal plant species, which brings about small-scale spatio-temporal changes in species abundance and hence species diversity.     

Competition between Shoots in Stands of Clonal Plants

Abstract Data on shoot turnover, shoot size distributions and individual shoot growth in stands of clonal plants are reviewed and compared with shoot density regulation in non-clonal plants. Shoots are the smallest units capable of independent existence in clonal as well as unitary plants and this comparison allows an investigation of the impact of integration and other factors on clonal shoot dynamics. Physiological integration, the capacity of clonal plants to exchange resources and growth substances between interconnected shoots, has long been held responsible for the ability of clonal plants to prevent shoot overproduction and for the general lack of density-dependent mortality and self-thinning. A review of published data leads to the conclusion that the effects of integration on shoot dynamics in clonal plants have been overestimated. Other characteristics such as the density-dependent control of meristem activity, the synchronization of shoot growth in spring and the maximum shoot size, seem more important for the deviating behaviour of shoot populations of clonal plants.     

Growth Dynamics of Shoot Height and Foliage Structure of a Rhizomatous Perennial Herb, Polygonum cuspidatum

The growth dynamics of shoot populations of Polygonum cuspidatumwere investigated at the Houei crater (approx. 2380 m abovesea level) on the south-eastern slope of Mount Fuji. At thisstudy site, a genetic individual of this species produces apopulation of shoots in the form of a patch occupying a certainground area. Generally, genetic individuals are located awayfrom each other and hence there is little interaction betweenindividuals. A large-sized individual occupying 31·2m² ground area with shoots, a medium-sized individual (5·6m²) and a small-sized individual (1·4 m²) were selectedfor this study. In each individual, growth was investigatedat the shoot level. The results were analysed based on the diffusionmodel. Early in the growing season in 1990, there was littledifference in LAI (leaf area index) and shoot density betweenthe individuals. Shoots of the small- and medium-sized individualsshowed size-independent height growth, whilst those of the larger-sizedindividual showed size-dependent height growth. Consequently,small-sized shoots of the small- and the medium-sized individualshad greater RGRs of shoot height growth than those of the largeindividual at the early stage. As a result, in the small- andmedium-sized individuals, cv (coefficient of variance) and skewnessof shoot height decreased with time. Increases in cv and skewnessof shoot height were found in the large-sized individual. Thesize-independent growth pattern of shoot height in the small-and medium-sized individuals during early growing stages isdifferent from the growth pattern of non-clonal plant species,in which plant height growth is positively size-dependent. Theexistence of a regulatory mechanism of shoot height growth issuggested for the small- and medium-sized individuals. The foliagestructure of the large-sized individual was different from thatof the medium- and small-sized individuals. The foliage structureof small- and medium-sized individuals was similar to the theoretical"optimal foliage structure" of plants. In clonal plant species,a genetic individual occupies a certain ground area with itsshoots. Therefore, "optimal foliage structure" per unit groundarea brings about maximization of photosynthetic rate for agenetic individual, which is consistent with the maximizationof fitness at the level of the individual plant.Copyright 1994,1999 Academic Press Clonal plant species, diffusion model, growth regulatory mechanism, genetic individual, maximization of fitness     

FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL HELIX ASPERSA MULLER

Feeding of the land snail Helix aspersa (Müller) was observedat monthly intervals. Three natural populations in Galicia (NW-Spain)were studied. At two sites only a few plants constituted thebulk of the diet and in spring the snails' diet had the highestdiversity (H'). In the third population feeding and distributionof Helix aspersa (Müller) were observed in a small plotwith permanent patches of Urtica dioica. Nearly one half offeeding snails fed upon Urtica dioica. Most of the other observations wereon Mentha suaveolens, Ranunculus repens and Gramineae. The diversity ofthe snails' diet showed seasonal variation with the maximumin the autumn months. Comparison between the availability ofthe different plant species and their contribution to the snails'diet showed that the snails did not eat at random; Urtica dioicawas eaten much more than expected from its occurrence and grasseswere strongly under-represented in the snails' diet. Temporalchanges of availability were significantly correlated with the amountseaten in the case of Urtica, but not for the other food plants.The distribution of the snails in the plot was significantlycorrelated with that of Urtica. Chemical analyses of the foodplants revealed Urtica as the species with the higher protein,ash and calcium contents. The strong preference of Helix aspersafor Urtica dioica could be explained by the value of Urticaas food or by its suitability as habitat for the snails. The largestproportions of green material in the snails' diet occurred inthe spring and juveniles ate more green material than adultsin the three populations. (Received 16 March 1998; accepted 30 November 1998)     

A Canopy Photosynthesis Model for the Dynamics of Size Structure and Self-thinning in Plant Populations

A dynamic model for growth and mortality of individual plantsin a stand was developed, based on the process of canopy photosynthesis,and assuming an allometric relationship between plant heightand weight, i.e. allocation growth pattern of plant height andstem diameter. Functions G(t, x), for the mean growth rate ofindividuals of size x at time t, and M(t,x), for the mortalityrate of individuals of size x at time t, were developed fromthis model and used in simulations. The dynamics of size structurewere simulated, combining the continuity equation model, a simpleversion of the diffusion model, with these functions. Simulationsreproduced several well-documented phenomena: (1) size variabilityin terms of coefficient of variation and skewness of plant weightincreases at first with stand development and then stabilisesor decreases with an onset of intensive self-thinning; (2) duringthe course of self-thinning, there is a power relationship betweendensity and biomass per unit ground area, irrespective of theinitial density and of the allocation-growth pattern in termsof the allometric parameter relating plant height and weight.The following were further shown by simulation: (a) competitionbetween individuals in a crowded stand is never completely one-sidedbut always asymmetrically two-sided, even though competitionis only for light; (b) plants of ‘height-growth’type exhibit a greater asymmetry in competition than plantsof ‘diameter-growth’ type, (c) the effect of competitionon the growth of individuals in a crowded stand converges toa stationary state, even when the stand structure still changesgreatly. All of these theoretical results can explain recentempirical results obtained from several natural plant communities.Finally, a new, general functional form for G(t, x) in a crowdedstand is proposed based on these theoretical results, insteadof a priori or empirical growth and competition functions. Canopy photosynthesis, competition mode, continuity equation, self-thinning, simulation, size distribution     

Effects of self-thinning of shoots on the nutrient budgets of Zizania latifolia

This experimental study focused on the seasonal changes and mobilization of nutrients between plant parts to understand the implications of self-thinning in the ecology of Zizania latifolia (Griseb.) Turcz. ex Stapf. The observations of shoot density, above- and belowground biomass, and total nitrogen and total phosphorus concentrations were conducted from February 2002 to August 2003. The biomass of shoots that died during the period and daily uptake of nutrients were determined. The shoot density sharply increased until mid-April and thereafter decreased significantly due to self-thinning of shoots. Total nitrogen and phosphorus concentrations of rhizomes decreased initially due to translocation to new shoots; however, the nutrients of rhizomes were slightly replenished from dead shoots during self-thinning. In contrast to other species, self-thinning of Z. latifolia shoots reallocates some of the minerals contained in the dead shoots back to the rhizomes, which can be regarded as a strategy to replenish the reduced resources of the rhizomes. The initial intensive growth of shoots can be regarded as a strategy to maintain resource competition.     

A Model for Growth and Self-thinning in Even-aged Monocultures of Plants

A theoretical model is derived from simple postulates to describethe rates of growth and mortality of plants in populations ofdifferent densities. The growth rate is described by a modificationof the logistic growth differential equation in which the increasein weight of an individual plant depends on its area, s_i ratherthan on its weight. The effective area for growth of a plantis reduced by an empirical function, f(s_i) with two terms: oneterm expresses the constraint imposed upon the increasing totalarea of plants by the limited physical area of the plot; theother term allows for a competitive advantage or disadvantagefor plants of varying sizes. Depending on the value of the parametercontrolling the relative competitive advantage term, intrinsicvariability between plants can be amplified or suppressed. Anindividual plant dies if the f(s_i) results in a negative growthrate for that plant. Computer simulations of the growth andsurvival of plants at different population densities were run.The results exhibit characteristics that appear realistic uponcomparison with published data: a survival of the fittest occurringduring thinning; a line of slope close to –3/2 boundingthe graphs of log weight versus log density; and the occurrenceof bimodality, associated with subsequent mortality, on frequencydistribution of log weight. computer logistic model, growth differential equation, density-effect, competition, mortality, self-thinning     

A Model for Mortality in a Self-thinning Plant Population

A model for mortality process in a self-thinning plant populationis proposed. It considers the spacial process but does not requirepositional information of each individual plant due to the assumptionsthat plants with interacting neighbours all greater than themselvesare the first to die and neighbours' sizes are mutually independentat each growth stage. Mortality of plants of size x at age t,M(t, x), is given as M(t, x) = m{P(t, x)}ⁿ where P(t, x) isthe proportion of plants of size greater than x at age t, andm and n are parameters. This model fits data from an experimentalplantation of Abies sachalinensis and will be useful for furtherdevelopment of the theoretical study of plant population growth. Abies sachalinensis Fr. Schm., self-thinning, mortality, size distribution, neighbourhood effect, spacial process model     

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.     

Stand Density Effects on Branching in an Annual Legume (Senna obtusifolia)

Effects of proximity to nearest neighbours on shoot morphologyand branch growth were examined for Senna obtusifolia (L.) I.& B., an annual legume species that displays wide variationin branching. Periodic surveys described location and type ofgrowth at all nodes on plants in regularly spaced monocultures.Stands with interplant distances of 15-50 cm (51-5 plants m^-2)formed closed canopies with similar amounts of leaf area andbiomass. Number of lower primary branches, their degree of curvature,and location of branch apices relative to the main stem respondedto interplant distance relatively early in shoot growth, beforecanopy closure. The final effects of ten-fold differences inplanting density on the formation of these ascending lower brancheswere only two-fold differences in number of vertical shoot axeswithin the stand. An additional study examined response of lowerbranches to an adjacent gap in the stand. Initial number anddirection of branch growth were not affected by location ofthe gap. The first effect of crowding by neighbours was on number ofbranches initiated by developing shoots. Differences in branchingsubsequently increased through differentiaI survival and directionof branch growth. Such alterations principally affected lateralexpansion of individual shoots, with little effect on radialsymmetry or vertical distribution of leaves.Copyright 1994,1999 Academic Press Branch morphology, canopy, competition, light quality, Senna obtusifolia, shoot growth, sicklepod     

Differences in Architecture and Shoot Growth during Stagnant and Extension Growth Phases of Acanthopanax sciadophylloides(Araliaceae)

The shoot growth of a deciduous tree, Acanthopanax sciadophylloidesFranch. et Savat. shows inter-annual intermittent repetitionof two distinctive phases, a stagnant growth phase (S-phase)and vigorous extension-growth (E-phase). To help understandthe differentiation mechanism, shoot development was studiedover time in both shoot phases. S-phase and E-phase shoots weredistinguished from each other by morphological traits: S-phaseshoots are characterized by higher allocation to leaves anda shorter period of stem growth, while E-phase shoots show continuousstem extension over the growing season. Specific leaf area didnot differ between the two phases. This shoot differentiationwas similar to the morphological differentiation of shoots betweenlong vs. short shoots found in some temperate trees. Leavesof both phases were well-dispersed through adjustment of petiolelength and leaf-blade size to reduce mutual shading within ashoot. Stem-wood density of current-year shoots was lower inE-phase compared with S-phase shoots. Leaves produced earlyin the season affected the growth phase of the following year.These results suggest that annual shoot differentiation of A.sciadophylloides was determined during the previous season andreflects leaf productivity in a given habitat during that growingseason. Copyright 2001 Annals of Botany Company Acanthopanax sciadophylloides, Araliaceae, biomass allocation, intermittent shoot growth, leaf display, shoot architecture, shoot differentiation     

Partitioning of Stored Resources Between Shoots in a Clone, and its Effects on Shoot Size Hierarchy

A theoretical study is described of the effects of the patternof partitioning of stored resources between the shoots of aclone on the development of shoot size hierarchy. When thereis an increasing convex relationship between the amount of resourcesavailable to a shoot at the start of growth and its biomassat maturity, the mode of competition between the shoots is asymmetricand the sharing of stored resources between shoots will notmaximize above-ground biomass of the clone at maturity. Underthis condition, the plant is predicted to allocate a smalleramount of its resources to storage, producing a lower below/above-groundbiomass ratio. Clonal species in which shoots compete asymmetricallyare known to have smaller below/above-ground biomass ratios.When there is an increasing concave relationship between theamount of resources available to a shoot at the start of growthand shoot biomass at maturity, the mode of competition willbe symmetric, and equal sharing of stored resources betweenall shoots at the start of growth will maximize the above-groundbiomass of the clone at maturity. If sharing of resources isthe optimal pattern of partitioning, all shoots of the cloneare predicted to be equal in size at maturity and, therefore,a size hierarchy will not develop within a clone. Copyright2001 Annals of Botany Company Clonal plants, pattern of partitioning, stored resources, shoot competition, size hierarchy, sharing of resources, symmetric competition     

Assessing Costs of Reproduction in Mountain Birch: The Importance of Considering the Modular Level

We studied the effects of production of male catkins on growthand the subsequent year's male catkin production in mountainbirch, Betula pubescens subsp. czerepanovii at both the shootand branch level. (Shoot is defined here as the product of asingle growing season while branch refers to a structure consistingof several shoots.) Like heterophyllous trees in general, mountainbirch canopy expansion takes place via the production of longshoots. We found phenotypic trade-offs between long shoot growthand male reproduction at the shoot level in the year of catkinproduction. Generative parental long shoots (long shoots withmale catkins) were significantly shorter than the vegetativeones (long shoots without a male catkin). In contrast, we foundno effects of male reproduction on the subsequent year's malecatkin production at the shoot level. Although the mean lengthof secondary long shoots (long shoots growing from the lateralbuds of parental long shoots) did not differ between vegetativeand generative parental long shoots, there was considerablebetween-individual variation in the response of individual trees.In addition, production of male catkins diminishes canopy expansionin mountain birch because the number of secondary long shootsproduced by generative parental long shoots was smaller thanthat of vegetative parental long shoots. At the branch level,the association between total long shoot growth and male catkinproduction was positive, i.e. no trade-off was found. This maybe because the strong sink strength (the ability of a branchto import assimilates from elsewhere in the tree) of branchesbearing reproductive long shoots masks possible trade-offs.We emphasize the importance of considering several levels ofthe modular hierarchy when analysing costs of reproduction inmodular organisms. Copyright 2000 Annals of Botany Company Hierarchical structure, modularity, mountain birch, short and long shoots, trade-off, Betula pubescens subsp.czerepanovii (Orlova) Hämet-Ahti     

A New Method for the Experimental Droughting of Plants

A new method of subjecting herbaceous plants to drought involvingdaily transfers of root systems between nutrient solutions andair is proposed. It appears to have the following advantagesover previous approaches: (1) standardization of the intensityand timing of drought; (2) avoidance of the confounding influenceof mineral nutrient stress; (3) allowance for a diurnal droughtingcycle; (4) provision for progressive cycles of drought; (5)complete retrieval of roots and shed plant parts. Allocation, drought, growth, moisture, stress, Urtica dioica     

Effects of Density and Extinction Coefficient on Size Variability in Plant Populations

The effects of density and extinction coefficient on size variability,as measured by the coefficient of variation of plant weightin even-aged monocultures, were investigated theoretically usinga diffusion model of growth and size distribution and a canopyphotosynthesis model over the range of densities at which self-thinning(size-dependent mortality) does not occur. Size inequality (thecoefficient of variation of plant weight) increases with increasingdensity or leaf area index at each growth stage. Plants witherect leaves are prone to lower size inequality than plantswith horizontal leaves. These results agree well with existingobservations on even-aged plant monocultures and suggest thatcompetition between plants is mainly one-sided (competitionfor light). One sided competition affects size variability througha G(t, x) function (mean growth of plants of size x at timet per unit time). Two-sided competition (including competitionfor nutrients) affects size variability through a D(t, x) function(variance of growth of plants of size x at time t per unit time).In this case, size inequality decreases with increasing density.The importance of studying size variability is emphasized. Helianthus annus L., size variability, size inequality, coefficient of variation, competition, density effect, extinction coefficient, diffusion model, canopy photosynthesis model     

Effect of Localized Placement of Nutrients on Root Competition in Self-thinning Populations

The hypothesis that increased root competition can lower theslope and/or intercept of the self-thinning line traversed byplant populations was tested using localized placement of nutrientsto increase root competition. Localized placement of nutrientswill result in increased root competition, if average inter-rootdistances are reduced, and if nutrients are in limiting supply.It was predicted that high-density, nutrient-limited populationsof Ocimum basilicum L. grown with localized placement of nutrientswould self-thin along a lower biomass–density line thannon-localized controls. This was tested at two fertility levelson a soil-based potting medium in expt 1, and at one fertilitylevel on washed sand in expt 2. Localized placement of nutrients significantly reduced the elevation(intercept) of the self-thinning line for both shoot and rootbiomass in expt 2. In expt 1, at the higher-fertility level,localized placement of nutrients had no significant effect;at the lower fertility level, localization had no significanteffect on thinning lines for shoot biomass, and resulted ina zero slope of the thinning line for root biomass. Canopy-based models of self-thinning failed to account for thereduction in the thinning-line intercept observed in expt 2.In both experiments, localized placement of nutrients resultedin a higher proportion of total root length being located inthe localization zone, which would result in a reduction inthe average inter-root distance. This would intensify root competitionunder conditions of nutrient limitation. The hypothesis thatintensified root competition would lower the self-thinning lineis supported by the results of expt 2. Localized placement of nutrients; root competition; shoot competition; root–shoot allocation; self-thinning; Ocimum basilicum ; sweet basil     

Structure and Dynamics of Populations of Japanese Barberry (Berberis Thunbergii DC.) in Deciduous Forests of New Jersey

Japanese barberry, Berberis thunbergii DC., has become a prominent exotic species in deciduous forests throughout the eastern and midwestern US. Populations range from small plants occurring at low densities to dense, impenetrable thickets of plants with up to 40 stems/individual. A study was undertaken at Morristown National Historical Park in New Jersey to document plant densities, plant size, recruitment through vegetative processes of new shoot initiation and clonal spread and recruitment from seedling establishment, and mortality of stems and plants. Nearly 2000 shoots on 370 plants were individually marked and followed for two growing seasons, and over 1000 seedlings were also individually marked and followed. Populations vary much more in total shoots/area than they do in plant individuals/area, or in mean plant size (shoots/plant), as even the sparse populations have a few large individuals. Shoot mortality is less than new shoot initiation, but most plants do not change in size or change by small numbers of stems. However, the number of new shoots per plant increases as plant size increases. Once plants have three stems, they suffer little or no mortality. Seedling establishment is proportional to the density of shoots, so that as plants grow in size, local recruitment from seed increases. Large numbers of seedlings, and a survival rate of 10%, combine to make seedling recruitment a major component of population increase. The combination of multiple forms of vegetative and seed-based population growth, and the very low rates of plant mortality due to the multi-stemmed growth form explains the ability of this invasive species to rapidly produce dense, persistent populations.     

Influence of plant maturity, shoot reproduction and sex on vegetative growth in the dioecious plant Urtica dioica

Background and Aims

Stinging nettle (Urtica dioica) is a herbaceous, dioecious perennial that is widely distributed around the world, reproduces both sexually and asexually, and is characterized by rapid growth. This work was aimed at evaluating the effects of plant maturity, shoot reproduction and sex on the growth of leaves and shoots.

Methods

Growth rates of apical shoots, together with foliar levels of phytohormones (cytokinins, auxins, absicisic acid, jasmonic acid and salicylic acid) and other indicators of leaf physiology (water contents, photosynthetic pigments, α-tocopherol and F_v/F_m ratios) were measured in juvenile and mature plants, with a distinction made between reproductive and non-reproductive shoots in both males and females. Vegetative growth rates were not only evaluated in field-grown plants, but also in cuttings obtained from these plants. All measurements were performed during an active vegetative growth phase in autumn, a few months after mature plants reproduced during spring and summer.

Key Results

Vegetative growth rates in mature plants were drastically reduced compared with juvenile ones (48 % and 78 % for number of leaves and leaf biomass produced per day, respectively), which was associated with a loss of photosynthetic pigments (up to 24 % and 48 % for chlorophylls and carotenoids, respectively) and increases of α-tocopherol (up to 2·7-fold), while endogenous levels of phytohormones did not differ between mature and juvenile plants. Reductions in vegetative growth were particularly evident in reproductive shoots of mature plants, and occurred similarly in both males and females.

Conclusions

It is concluded that (a) plant maturity reduces vegetative growth in U. dioica, (b) effects of plant maturity are evident both in reproductive and non-reproductive shoots, but particularly in the former, and (c) these changes occur similarly in both male and female plants.     

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.