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.     

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.     

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.     

Glandular Hairs and Essential Oil in Developing Leaves of Ocimum basilicum L. (Lamiaceae)

The types of glandular hairs and their pattern of distributionon leaves of Ocimum basilicum L. at different stages, were investigated.Their essential oil at the various stages was chemically analysed.The density of the glandular hairs appears to be very high onyoung meristematic leaves and on meristematic regions of olderleaves. When the leaf regions start to expand no new glandularhairs are produced and their density decreases. The essentialoil differs in the percentage of some of its components betweenyoung and mature leaf regions.Copyright 1993, 1999 AcademicPress Ocimum basilicum, sweet basil, glandular hairs, leaf development, essential oil, cugenol, methyl chavicol, 1,8-cineole     

Shoot Growth and Mortality Patterns of Urtica dioica, a Clonal Forb

The growth and mortality patterns of the clonal forb Urticadioica were investigated at the level of the individual shootin two growing seasons, 1991 and 1992, in a natural stand. Shootheight and diameter at ground level of each shoot tagged inspring were measured repeatedly five times during the growingseason. Dry weights of these repeatedly measured shoots wereestimated using an allometric relationship between dry weight,height and diameter of harvested shoots. A large decrease inshoot density occurred with stand development from the beginningof the growing season in both the years: (1) shoot survivalrate was about 30% at the end of the growing season; (2) shootmortality rate per 10 x 10 cm subplot between censuses was positivelydependent on shoot density per subplot; (3) the mortality rateof individual shoots was negatively dependent on shoot size(height, diameter and weight) at each growing stage, suggestingone-sided competition between living and dying shoots; (4) shootsize (height, diameter and weight) variability in terms of thecoefficient of variation and skewness decreased in accordancewith shoot mortality. Symmetric competition between living shootswas detected by regression analysis based on a model for individualshoot growth considering the degree of competitive asymmetry.However, the competitive effect on individual shoot growth wasvery small (nearly absent). The mortality pattern of Urticadioica indicates that shoot self-thinning occurred from theearly growing stage as in non-clonal crowded monospecific stands,and contrasts with many clonal plants where shoot self-thinningrarely occurs or, if any, is confined only to a short periodof the later growing stage. The pattern of growth and competitionbetween living shoots of Urtica dioica contrasts with non-clonalcrowded plants undergoing intense competition (usually asymmetric)between individuals, but is a common feature of many clonalplants where shoot competition is supposed to be reduced by'physiological integration' between shoots. These form a newpattern not reported yet for clonal plants. It is pointed outthat clonal plants show a wider spectrum of the growth, competitionand mortality patterns of shoots than non-clonals. Some possiblemechanisms for the pattern of Urtica dioica are discussed.Copyright1995, 1999 Academic Press Shoot competition, diffusion model, individual shoot growth, shoot self-thinning, shoot size variability, Urtica dioica L     

Yield-density Relationships: the Influence of Resource Availability on Growth and Self-thinning in Populations of Vulpia fasciculata

Monocultures of Vulpia fasciculata were grown over a wide rangeof densities to investigate the influence of crowding and nutrientsupply on growth and self-thinning. For a given time and densityseries the relationship between mean yield per plant (w) andthe density of survivors (N) could be described by the equation w= w_m (1+aN)^–b. where w_m is the yield of an isolated plant, a is the area requiredto achieve a yield of w_m and b describes the effectiveness withwhich resources are taken up from the area. All three parametersincreased with time. Adding nutrients changed not only the rate at which the effectsof crowding occurred but also the intensity of crowding since w_m = C(a^b)^D. where C and D are constants. The addition of nutrients resultedin an increase in the value of C. Such an increase means thata larger weight can be supported by a given area because theresources within that area are greater. During the early phases of growth, populations of V. fasciculataconformed to the –3/2 power law, w = cN^–3/2, butonly at very high densities with a plentiful supply of nutrients.However, once the maximum standing crop had been reached thetrajectory of the thinning line switched to a slope of justless than –1 when weight was ploted against density onlogarithmic scales. The intercept of the –3/2 thinningline was considerably higher (log c = 5.74) than those for mosttrees and forbs but was similar to those of a number of othergrasses. Vulpia fasciculata, dune fescue, yield-density models, self-thinning, density-dependence, nutrient supply     

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.     

Facilitation among plants can accelerate density‐dependent mortality and steepen self‐thinning lines in stressful environments

The speed and slope of plant self‐thinning are all affected by plant–plant interactions across environmental gradients. Possible mechanisms driving the self‐thinning dynamics include the relative strength of root versus shoot competition, and the interplay between competition and facilitation. Although these mechanisms often act in concert, their relative importance has not yet been fully explored. We used both a one‐layer and a two‐layer zone‐of‐influence (ZOI) model to examine how competition and facilitation drive self‐thinning across stress gradients. As a development of the traditional ZOI model, the two‐layer version explicitly models shoot and root growth and neighbor interactions, and thus the overall size‐symmetry of competition is regulated by the relative strength of root versus shoot competition. One‐layer model simulations revealed that increasingly asymmetric competition accelerated thinning, and steepened (slope ranged from about –1 to –4/3) and lowered self‐thinning lines. Stress slowed down density‐dependent mortality considerably when competition was not completely symmetric. Stress significantly decreased the self‐thinning intercept, while facilitation simply counteracted stress effects. Both stress and facilitation showed little effect on the slope. In the two‐layer model, both stress and facilitation affected mortality in the same way as in the one‐layer version when competition was not completely symmetric. Different from the one‐layer model, the two‐layer version showed that the effects of stress and facilitation on the self‐thinning slope were mediated by the asymmetry of competition. As stress increased, the overall asymmetry of competition shifted from asymmetric to symmetric due to increased relative strength of root competition. High stress thus dramatically flattened self‐thinning lines, whereas the inclusion of facilitation counteracted stress and led to steeper self‐thinning lines. Our two‐layer model is based on the current knowledge of plant–plant interactions, and better represents ecological realities. It can help elaborate experiments for testing the role of competition and facilitation in driving plant population dynamics.     

Relationship between the virtual dynamic thinning line and the self-thinning boundary line in simulated plant populations

The self-thinning rule defines a straight upper boundary line on log-log scales for all possible combinations of mean individual biomass and density in plant populations. Recently, the traditional slope of the upper boundary line, -3/2, has been challenged by -4/3 which is deduced from some new mechanical theories, like the metabolic theory. More experimental or field studies should be carried out to identify the more accurate self-thinning exponent. But it＇s hard to obtain the accurate self-thinning exponent by fitting to data points directly because of the intrinsic problem of subjectivity in data selection. The virtual dynamic thinning line is derived from the competition-density （C-D） effect as the initial density tends to be positive infinity, avoiding the data selection process. The purpose of this study was to study the relationship between the virtual dynamic thinning line and the upper boundary line in simulated plant stands. Our research showed that the upper boundary line and the virtual dynamic thinning line were both straight lines on log-log scales. The slopes were almost the same value with only a very little difference of 0.059, and the intercept of the upper boundary line was a little larger than that of the virtual dynamic thinning line. As initial size and spatial distribution patterns became more uniform, the virtual dynamic thinning line was more similar to the upper boundary line. This implies that, given appropriate parameters, the virtual dynamic thinning line may be used as the upper boundary line in simulated plant stands.     

Alginate encapsulation of axillary buds of Ocimum americanum L. (hoary basil), O. Basilicum L. (sweet basil), O. Gratissimum L. (shrubby basil), and O. Sanctum. L. (sacred basil)

Summary Propagation and conservation of four pharmaceutically important herbs, Ocimum americanum L. syn. O. canum Sims. (hoary basil); O basilicum L. (swett basil); O. gratissimum L. (shrubby basil); and O. sanctum L. (sacred basil) was attempted using synthetic seed technology. Synthetic seeds were produced by encapsulating axillary vegetative buds harvested from garden-grown plants of these four Ocimum species in calcium alginate gel. The gel contained Murashige and Skoog (MS) nutrients and 1.1-4.4 μM benzyladenine (BA). Shoots emerged from the encapsulated buds on all six planting media tested. However, the highest frequency shoot emergence and maximum number of shoots per bud were recorded on media containing BA. Of the six planting media tested, both shoot and root emergence from the encapsulated buds in a single step was recorded on growth regulator-free MS medium as well as on vermi-compost moistened with halfstrength MS medium. Rooted shoots were retrieved from the encapsulated buds of O. americanum, O. basilicum, and O. sanctum on these two media, whereas shoots of O. gratissimum failed to root. The encapsulated buds could be stored for 60 d at 4°C. Plants retrieved from the encapsulated buds were hardened off and established in soil.     

Effect of Water Deficit on Self-thinning Line in Spring Wheat （Triticum aestivum L.） Populations

Monocultures of spring wheat （Triticum aestivum L.） were grown at overcrowded densities （10 000 and 3 000 plants per m^2） under well-watered and water-stressed conditions to investigate the effects of water deficits on self-thinning. The results showed that density reduction in water-stressed populations was delayed compared with that In well-watered populations. Populations grown In well-watered conditions conformed to the -3/2- power law. Compared with the well-watered condition, there was no significant decrease of the self-thinning line under water-stressed conditions In this experiment, although the rate of average shoot blomass accumulatlon decreased. This result Implied that the exponent of the -3/2-power equation Is not as sensitive as the rate of average shoot blomass accumulation to water stress. Further analysis indicated that, In each density treatment, the lines of the height versus shoot blomass relationships did not differ significantly between the two water conditions. However, the Intercepts of the height versus shoot blomass relationships were greater In the higher-density populations （10 000/m^2） than those In the lower-density populations （3 000/m^2）. These results showed that water deficit did not change plant geometry In this experiment. That Is to say, shoot competition for light remains constant at a given blomass, although root competition for water becomes more serious In water deficit conditions. Based on these results and previous reports we propose that, to affect the thinning line slope, changes In symmetric competition are not as efficient as changes In asymmetric competition.     

In Defence of the -3/2 Boundary Rule: a Re-evaluation of Self-thinning Concepts and Status

The -3/2 power rule, or -3/2 self-thinning rule, was accepted10 years ago as an important generalization, but has recentlybeen questioned by a number of authors. This paper assesseswhat remains of the rule. While it has been empirically establishedthat size-density trajectories followed by self-thinning plantpopulations do not necessarily follow a -3/2 slope, a more generalpower rule describing a density-dependent upper limit to meanshoot biomass per plant (the '-3/2 boundary rule') remains largelyintact. Principal component analysis (PCA) overestimates the steepnessof the thinning slope if y:x variance ratio is greater than1:1. Lonsdale's (Ecology 71: 1373-1388) overall mean PCA slopeof -0·6 for biomass-density suggests a true mean slopeclose to the theoretical value of -0·5. Reduced majoraxis (RMA) regression appears a reasonable approximation forthe -3/2 but not the -1/2 formulation of the rule. Fitting ofa linear functional relationship (LFR) is a more appropriateslope estimation procedure, not previously used for data onthinning. None of these procedures estimates a boundary linethat is not transgressed by any data point except through errorsof measurement. Mortality due to overcrowding ensues when a small, suppressedplant no longer holds its leaves high enough in the canopy tomaintain a positive carbon balance. It follows that LAI shouldremain constant during thinning, and that self-thinning theoryshould be developed in terms of maximum leaf area index andthe biomass required to support it. A derivation is presentedand some of its consequences are examined.Copyright 1995, 1999Academic Press Self-thinning, -3/2 power rule, -3/2 self-thinning rule, boundary line, size-density compensation, regression methods     

The Influence of Mineral Nutrition on the Root Development of Trees: I. THE GROWTH OF SITKA SPRUCE WITH DIVIDED ROOT SYSTEMS

Sitka spruce seedlings were grown with their root systems dividedbetween two contrasting nutrient regimes. One half of the rootsystem was supplied with a solution containing N, P, and K ata range of concentrations while the untreated half receivedwater only. High-nutrient treatments induced two flushes ofshoot growth resulting in a large shoot system, whereas plantsin the low-nutrient treatments flushed once only and showedsymptoms of nutrient deficiency. Root growth, assessed in terms of dry weight and diameter ofboth primary and woody tissues, was stimulated in the rootsto which the nutrients were actually applied, whereas in theuntreated roots on treated plants only the primary root diameterwas enhanced. However, internal nutrient concentrations on bothsides of the root system were related to the concentrationsapplied, though to a slightly lesser extent in the untreatedroots. Thus, the nutrients which had been internally translocatedto the untreated roots had little effect on their growth. The localized stimulation of xylem production in the woody rootsextended into the stem along a spiral pathway which was demonstratedby the movement of dye. Possible mechanisms are discussed by which differential rootgrowth is brought about by a localized supply of mineral nutrients.     

The Nitrogen Content of Plants and the Self-thinning Rule of Plant Ecology: A Test of the Core-skin Hypothesis

The ‘core-skin’ hypothesis postulates that secondarilythickened plants behave energetically as an inert ‘core’covered by an active ‘skin’, the ‘skin’being two-imensional, the ‘core’ three-dimensional.This would explain the ‘self-thinning ‘or‘–3/2’ rule of plant ecology, that is, the tendencyfor log (dry weight per plant) and log (number of plants perunit area) to progress along a straight line relationship, withslope = – 3/2’. The hypothesis was tested as follows. Plant nitrogen contentwas used as an estimate of the mass of ‘skin’ perplant, and dry weight as an estimate of the mass of the ‘core’.As plants mature the slope of the relationship between y = log(mass of nitrogen per plant) and x = log (mass of dry matterper plant) is expected to decline from an initial value of 1.0towards a final value of 0.66. The intercept of the relationshipis expected to reflect the intrinsic content of ‘skin’per unit of ‘core’. Genotypic variation in thisparameter should cause genotypic differences in the maximumattainable yield of biomass per unit area. The expectations were investigated by fitting the function y= p+qx+r exp – x to 30 sets of data on plant nitrogencontent, plant weight and time in 18 different vegetables. Simplelinear regressions of y on x were fitted to more limited setsof data on weights and nitrogen contents of mature trees. Theexpectations were, with some minor exceptions, confirmed. Nitrogen, yield, plant competition, self-thinning     

Effect of Root and Shoot Meristem Temperature on Shoot to Root Dry Matter Partitioning and the Internal Concentrations of Nitrogen and Carbohydrates in Maize and Wheat

Maize (Zea mays L.) and spring wheat (Triticum aestivum L.)were grown in nutrient solution at uniformly high air temperature(20 °C), but different root zone temperatures (RZT 20, 16,12 °C). To manipulate the ratio of shoot activity to rootactivity, the plants were grown with their shoot base includingthe apical meristem either above (i.e. at 20 °C) or withinthe nutrient solution (i.e. at 20, 16 or 12 °C). In wheat, the ratio of shoot:root dry matter partitioning decreasedat low RZT, whereas the opposite was true for maize. In bothspecies, dry matter partitioning to the shoot was one-sidedlyincreased when the shoot base temperature, and thus shoot activity,were increased at low RZT. The concentrations of non-structuralcarbohydrates (NSC) in the shoots and roots were higher at lowin comparison to high RZT in both species, irrespective of theshoot base temperature. The concentrations of nitrogen (N) inthe shoot and root fresh matter also increased at low RZT withthe exception of maize grown at 12 °C RZT and 20 °Cshoot base temperature. The ratio of NSC:N was increased inboth species at low RZT. However this ratio was negatively correlatedwith the ratio of shoot:root dry matter partitioning in wheat,but positively correlated in maize. It is suggested that dry matter partitioning between shoot androots at low RZT is not causally related to the internal nitrogenor carbohydrate status of the plants. Furthermore, balancedactivity between shoot and roots is maintained by adaptationsin specific shoot and root activity, rather than by an alteredratio of biomass allocation between shoot and roots.Copyright1994, 1999 Academic Press Wheat, Triticum aestivum, maize, Zea mays, root temperature, shoot meristem temperature, biomass allocation, shoot:root ratio, carbohydrate status, nitrogen status, functional equilibrium     

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.     

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     

The Consequence of Selection for Copper Tolerance on the Uptake and Accumulation of Copper inMimulus guttatus

Previous research has shown that copper tolerance inMimulusguttatusFischer ex DC. is controlled by a single major geneand can be enhanced by a number of minor genes (or modifiers).Here we report the uptake of copper by three lines which allpossessed the major tolerance gene but differed in the modifiergenes: the major gene only (IT), the major gene plus increased(HT6) and decreased (LT6) modifiers. HT6 showed the highestcopper tolerance and IT the lowest. Copper uptake was investigatedat five copper concentrations and over 30 d to analyse the concentrationof copper accumulated in roots and shoots and the partitioningof apoplastic and symplastic root copper. Significant differenceswere found for root copper concentration with the IT line accumulatingthe highest levels. Fifty-two per cent of the root copper inthe IT line is symplastic and this increases to 60% in LT6 and64% in HT6. Significant differences were recorded for shootcopper concentration with HT6 accumulating the highest and ITthe lowest. At the highest external copper concentration theHT line accumulated nearly 800 µg g^-1in its shoot, approachinglevels reported for copper hyperaccumulation.Copyright 1997Annals of Botany Company Copper tolerance; copper uptake; copper minor genes; Mimulus guttatus     

The Effect of Sugar Supply Rate on Photosynthetic Development of Ocimum basilicum (Sweet Basil) Cells in Continuous Culture

The fructose supply rate to continuous cultures of Ocimum basilicumL. cells was changed in two ways. The concentration of fructosein the medium feed was diminished or the dilution rate was increasedin fructose-limiting conditions. When fructose in the feed wasdecreased, whilst maintaining constant dilution rate (0.87 µ_max),the specific rate of chlorophyll production increased—thebiomass became greener. Actual photosynthesis rate (dry biomass^–1)measured in the steady state conditions also increased. When fructose supply rate was increased by increasing the dilutionrate, the specific rate of chlorophyll production and actualphotosynthesis rate (dry biomass^–1) increased until acritical dilution rate (0.64 µ_max) was reached, thereafterdecreasing. The potential photosynthesis rate was measured on samples ofcells supplied with additional photons. This was inversely relatedto dilution rate. It seemed possible that the concentration of residual fructoseoutside the cells was related to specific growth rate but probablynot according to the Monod model. The specific production ofchlorophyll may be regulated by the intracellular concentrationof a catabolite of fructose, possibly glucose. Results suggestthat the specific production rate of chlorophyll was inhibitedwhen glucose concentration in the cells was above a thresholdof about 1.2% dry biomass. The degree of inhibition was a functionof glucose concentration above this threshold. Key words: Continuous culture, Photosynthesis, Greening, Ocimum basilicum     

Effects of competition on root–shoot allocation in <Emphasis Type="Italic">Plantago lanceolata</Emphasis> L.: adaptive plasticity or ontogenetic drift?

We investigated how shoot and root allocation in plants responds to increasing levels of competitive stress at different levels of soil fertility. In addition, we analyzed whether different responses were due to adaptive plasticity or should be attributed to ontogenetic drift. Plantago lanceolata plants were grown during 18 weeks at five plant densities and four nutrient supply levels in pots in the greenhouse. Thereafter root and shoot biomass was measured. There were clear negative effects of increasing plant densities on plant weights revealing strong intraspecific competition. At the lower N-treatments, the proportional allocation to root mass increased with increasing competitive stress, indicating the important role of belowground competition. At the higher N-supply rate, the relationship between competitive stress and shoot to root ratio was neutral. These responses could not be attributed to ontogenetic drift, but could only be explained by assuming adaptive plasticity. It was concluded that at lower N-supplies belowground competition dominates and leads to increased allocation to roots, while at the higher N-supply competition for soil resources and light had balanced impacts on shoot and root allocation. An alternative hypothesis explaining the observed pattern is that light competition has far less pronounced impacts on root–shoot allocation than nutrient deprival.