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.     

Theoretical Relationships Between Mean Plant Size, Size Distribution and Self Thinning under One-sided Competition

As yet there is no comprehensive theory in plant populationecology to explain relationships between mean plant size, sizedistribution and self-thinning. In this paper, a new synthesisof plant monocultures is proposed. If the reciprocal relationshipbetween plant biomass and plant population density among variousstands of even-aged plant populations holds, the same reciprocalrelationship must exist between cumulative mass and cumulativenumber of plants from the largest individual within a population,assuming strict one-sided competition (which is an extreme conditionfor competition for light among plants). The two parametersof the relationship between cumulative mass and cumulative numberwithin a stand both correlate with maximum plant height in thestand. One parameter equals the reciprocal of the potentialmaximum plant mass per area, which is expressed by the productof maximum plant height and dry-matter density. The other parametercorrelates with the potential maximum individual plant mass,which is allometrically related to maximum plant height. Asa stand develops, the growth rate of the smallest individualswill become zero due to suppression from larger individuals,and they will die; i.e. self-thinning will occur. The slopeof the self-thinning line is expressed through the coefficientsof allometry between height and mass and between dry matterdensity and height. When the former coefficient is 3 and thelatter is 0, the gradient exactly corresponds to the value expectedfrom the 3/2 power rule, but it can take various values dependingon the values of the two coefficients. Competition among individualsdetermines size-density relationships among stands, which inturn determine the size structure of the stand. The size structureconstrains the growth of individuals and results in self-thinningwithin the stand.Copyright 1999 Annals of Botany Company. Monoculture, plant population, self-thinning, competition, hierarchy, size-structure.     

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

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

Effects of adaptation to winter stress on biomass production, growth and morphology of three contrasting white clover cultivars

Changes in biomass production and partitioning, together with morphology and growth of three white clover ( Trifolium repens L.) cultivars after exposure to winter stress were recorded in a controlled environment study. The cultivars used were Grasslands Huia (H, New Zealand), AberHerald (AH, bred in UK from material of Swiss origin) and Sandra (S, Sweden), sub-populations of these (survival populations), and a wild ecotype from the Uppsala area. The sub-populations were collected after 2 (with the exception of S) and 4 years of winter stress, respectively, at Uppsala (59°49' N, 17°39' E). In the controlled environment study the photoperiod was 12 h and there were two temperature regimes, namely 24/18 and 12/9°C (day/night), respectively. The experimental plants were cloned from eight different plants of each population. There were few interactions between populations and temperature regimes. Biomass production was lowest in the Swedish populations and in the surviving populations of the other cultivars collected after 4 years of exposure to winter stress. However, biomass partitioning, with the exception of allocation to branches, remained unaffected. Morphology differed significantly between cultivars, but, with the exception of internode length, was unaffected by exposure to winter stress. Internode length was significantly reduced in surviving populations of both H and AH after exposure to four winters, when its value was similar to that of the Swedish populations. The reduced internode length was a result of a greater reduction in the stolon extension rate than in the leaf appearance rate of survival populations compared with the original populations. It is hypothesised that reduced internode length would have implications for the carbon economy of the plant.     

Relationships Between Population Growth and Population Density in Monocultures of Larix leptolepis

One year-old Japanese larch (Larix leptolepis) seedlings were planted in a nursery from 1984 to 1986 at four density levels with four replicates in order to investigate the relationships between plant growth and density of survivors. As the results shown, self-thinning occurred severely in populations growing at high densities. The rate of the self-thinning followed a trajectory defined by the 3/2 power law. The relationship between mean dry weight per tree and population density at a given stage of growth followed the reciprocal equation and power equation at early experimental stages, but changed gradually to follow the 3/2 power law as plants grew further. Shch a change was mainly caused by the constant basal area per ms and decreased growth of tree height along a gradient of the density. The relationship between tree height (h) and density (p) also be discussed and in the 3/2 power law system it could be described by an equation consisting of h=αp（-1/2） where α is a constant.     

Ontogenetic changes in genetic variances of age-dependent plasticity along a latitudinal gradient

The expression of phenotypic plasticity may differ among life stages of the same organism. Age-dependent plasticity can be important for adaptation to heterogeneous environments, but this has only recently been recognized. Whether age-dependent plasticity is a common outcome of local adaptation and whether populations harbor genetic variation in this respect remains largely unknown. To answer these questions, we estimated levels of additive genetic variation in age-dependent plasticity in six species of damselflies sampled from 18 populations along a latitudinal gradient spanning 3600 km. We reared full sib larvae at three temperatures and estimated genetic variances in the height and slope of thermal reaction norms of body size at three points in time during ontogeny using random regression. Our data show that most populations harbor genetic variation in growth rate (reaction norm height) in all ontogenetic stages, but only some populations and ontogenetic stages were found to harbor genetic variation in thermal plasticity (reaction norm slope). Genetic variances in reaction norm height differed among species, while genetic variances in reaction norm slope differed among populations. The slope of the ontogenetic trend in genetic variances of both reaction norm height and slope increased with latitude. We propose that differences in genetic variances reflect temporal and spatial variation in the strength and direction of natural selection on growth trajectories and age-dependent plasticity. Selection on age-dependent plasticity may depend on the interaction between temperature seasonality and time constraints associated with variation in life history traits such as generation length.     

Combining winter hardiness and forage yield in white clover (Trifolium repens) cultivated in northern environments

Background and Aims

White clover (Trifolium repens) is an important component of sustainable livestock systems around the world. Its exploitation for agriculture in the northern, marginal areas is, however, currently limited by the lack of cultivars that combine persistence and high production potential. The aims are to investigate whether it is feasible to create breeding material of white clover for these areas by combining winter hardiness of northerly populations with good yielding ability of more southerly cultivars.

Methods

A total of 166 crosses of 14 different parental combinations between winter-hardy, low-yielding populations of northern origin and high-yielding commercial cultivars of more southerly origin were tested under field conditions in Iceland and Norway and the parental populations were compared in Norway. Spaced plants were transplanted into a smooth meadow grass (Poa pratensis) sward. Dry matter yield was estimated for 2 years after planting in Norway and morphological characters associated with yielding capacity were measured at both sites.

Key Results

The results showed that southerly cultivars had larger leaves and higher yielding potential than northern types but suffered more winter damage. Significant variation was found between full-sib families within the different parental combinations for all morphological characteristics measured in all three trials. However, it was difficult to detect any consistent morphological patterns between progeny groups across trial sites. No significant correlations were found between leaflet area and survival.

Conclusions

The present study has confirmed that it should be possible to simultaneously select for good winter survival and larger leaves and, hence, higher yielding ability under marginal conditions.Key words: Breeding, morphology, spaced plants, Trifolium repens, winter survival, progeny testing     

A model of physiological and allometric factors in the self-thinning curve

A generalized self-thinning curve for plants is derived from the modified Von Bertallanfy equation. When an asymptotic relation between photosynthesis per unit of leaf area and stocking density is assumed, the self-thinning curve thus derived is also asymptotic on a log-log scale but is fitted quite well by a log-linear approximation. The model predicts that the slope of the log-linear approximation is a function of (a) photosynthetic response to density and (b) the relation between leaf area and total aboveground biomass. Intercept of the log-linear approximation is a function of these plus maximum attainable biomass, site productivity, density at which maximum photosynthesis is attained, and the nature of carbon loss within the plant community. Linkages between various parameters within the model act to reduce differences in slope and intercept for species with different life history's and physiological requirements.     

Testing the cultivar vigor hypothesis: comparisons of the competitive ability of wild and cultivated populations of Pascopyrum smithii along a restoration chronosequence

Restoration practitioners often rely on seeds of widely available cultivars representing native species but nonlocal germplasm. Cultivation improves the supply of plant materials and minimizes revegetation costs, but can also favor agronomic traits, and resulting vigor may affect the competitive ability and long‐term persistence of cultivated genotypes at restoration sites. We compared cultivated, restored, and wild populations of Pascopyrum smithii (western wheatgrass) in a greenhouse study to test the extent to which cultivars outcompete local plants in biomass production, and to determine if morphological differences (including height and number of leaves) among cultivated and wild populations persist at restoration sites over time. We found evidence of vigor and greater competitive ability of cultivars in seed mass, growth rate, plant height, and biomass and this advantage occurred when plants were grown alone or in competition with other seed sources. Cultivar vigor persisted at restoration sites over 30 years, but restored populations more closely resembled wild, local populations when cultivars were planted in closer proximity to nearby undisturbed sites. This study supports the cultivar vigor hypothesis and provides evidence for the long‐term persistence of cultivated traits in the environment.     

AN EXPERIMENTAL EVALUATION OF DENSITY AND PLANT SIZE IN TWO LARGE BROWN SEAWEEDS

The effects of density on the growth rate and survival of individual plants as well as changes in population structure (hierarchy) and biomass accumulation (self-thinning) were experimentally evaluated in two brown macroalgae. Laminaria digitata (Hudson) Lamouroux and Fucus serratus Linnaeus populations were constructed at five (650–5156 plants·m^?2) and four densities (650–2668 plants·m^?2), respectively, and were cultivated in tanks. The relative growth rates and survivorship of individuals and the populations’ biomass and density (estimated dry mass) were periodically measured. To investigate how plant population size hierarchies influence conspecifics, single density populations of L. digitata were constructed of up to three sizes of plants in equal proportions, and these parts of the populations were compared through time with plants of the three sizes grown singly. At higher density, L. digitata plants grew more slowly, while F. serratus populations showed a similar trend that was never statistically significant. Survival of plants of both species was lower at high densities, and mortality selectively removed smaller plants. Plants of both species exhibited zero growth rates before death, when parts of the fronds were lost, but meristems (apical in F. serratus, at the base of the frond in L. digitata) were preserved until the death of the plant. All singly grown L. digitata plants survived, but survivorship was low in the fractions of small plants in mixed-size populations compared with that of the largest size plant fractions. Small L. digitata plants grew relatively faster than did large ones singly, but in mixed-size populations, small plants grew relatively slowly. Plant sizes became progressively more unequal (Gini and skewness coefficients) until self-thinning started reducing the size variability. The seaweeds followed self-thinning (density-biomass) trajectories predicted by the self-thinning “rule”, and self-thinning appeared to be seasonal- rather than species-dependent, as it occurred at a time of year when ambient light levels start to fall in the Isle of Man. Culture studies of this kind, despite their considerable potential, are a tool as yet underexploited by marine ecologists as a means of assessing intraspecific competitive interaction among seaweeds.     

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

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

Plant mortality varies with arbuscular mycorrhizal fungal species identities in a self-thinning population

Because arbuscular mycorrhizal fungal (AMF) species differ in stimulating the growth of particular host plant species, AMF species may vary in their effects on plant intra-specific competition and the self-thinning process. We tested this hypothesis using a microcosm experiment with Medicago sativa L. as a model plant population and four AMF species. Our results showed that the AMF species Glomus diaphanum stimulated host plant growth more than the other three AMF species did when the plants were grown individually. Glomus diaphanum also induced the highest rate of mortality in the self-thinning plant populations. We also found a positive correlation between mortality and growth response to colonization. Our results demonstrate that AMF species can affect plant mortality and the self-thinning process by affecting plant growth differently.     

On the Ordered Development of Plants 2. Self-thinning in Plant Communities

It is shown that the hypothesis that a growing point on a vegetativeplant requires a minimum rate of supply of assimilate to continuegrowth can quantitatively describe self-thinning in communitiesof Trifolium subterraneum. The hypothesis can also be used toexplain the different relationships observed between mean plantweight and plant density when plants are grown in full daylight,70 per cent shade, and transferred between the two light environments.The implications of the hypothesis to self-thinning in naturalplant communities are discussed. Self-thinning, assimilate, plant development     

Local habitat conditions explain the variation in the strength of self‐thinning in a stream salmonid

Self-thinning patterns are frequently used to describe density dependence in populations on timescales shorter than the organism''s life span and have been used to infer carrying capacity of the environment. Among mobile animals, this concept has been used to document density dependence in stream salmonids, which compete over access to food and space. The carrying capacity, growth conditions, and initial cohort sizes often vary between streams and stream sections, which would influence the onset and strength of the density dependence. Despite much effort in describing habitat relationships in stream fishes, few studies have explicitly tested how the physical environment affects the slope of the thinning curves. Here, we investigate the prevalence and strength of self-thinning in juvenile stages of a steelhead (Oncorhynchus mykiss) population in Idaho, USA. Further, we investigate the roles of local physical habitat and metabolic constraints in explaining the variation in thinning curves among study sites in the watershed. Only yearling steelhead exhibited an overall significant thinning trend, but the slope of the mass–density relationship (−0.53) was shallower than predicted by theory and reported from empirical studies. There was no detectable relationship in subyearling steelhead. Certain abiotic factors explained a relatively large portion of the variation in the strength of the self-thinning among the study reaches. For subyearling steelhead, the slopes were negatively associated with the average water depth and flow velocity in the study sites, whereas slopes in yearlings were steeper in sites that incurred a higher metabolic cost. Our results show that the prevalence and strength of density dependence in natural fish populations can vary across heterogeneous watersheds and can be more pronounced during certain stages of a species'' life history, and that environmental factors can mediate the extent to which density dependence is manifested in predictable ways.     

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

Background and Aims

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

Methods

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

Key Results

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

Conclusions

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

植物种群自疏过程中构件生物量与密度的关系

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

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.     

VARIATION AMONG POPULATIONS OF XANTHIUM STRUMARIUM (COMPOSITAE) FROM NATURAL AND RUDERAL HABITATS

We assessed the mechanisms underlying the ability of cocklebur to spread from its natural riverside habitats and establish weedy populations in urban waste areas. We collected fruits from plants growing in natural and urban ruderal habitats and planted 2 maternal families from each of 9 habitat populations in 3 experimental gardens. The gardens were all in full sunlight but differed in the availability of water and nutrient resources. Plant performance in the gardens was measured by numbers and size of fruits produced. Traits known to be associated with cocklebur reproductive success were also measured: times to emergence and anthesis, photosynthetic capacity, mean stomatal conductance, relative growth rate, and biomass allocation to leaves and stems. Although there were significant differences among populations in the tradeoff between rate of growth in height and timing of anthesis, these population differences were not associated with habitat. Apart from a tendency to produce larger fruits in ruderal populations, there were no detectable differences in the characteristics of plants from natural vs. ruderal habitats. Plants from both habitats did have substantial and significant plastic responses to growth environment. In the three experimental gardens, fruit numbers increased with resource availability but fruit size did not differ significantly. As resource availability increased, plants from both habitats sustained growth longer and became more branchy. Canonical discriminant analysis of all the somatic and reproductive traits together supported the idea that natural and ruderal populations do differ in their overall plastic response to resource availability. The subtly different plastic responses of plants from the two habitats do not arise by substantial adjustments in a few dominant traits, but instead by relatively minor adjustments in a host of functionally interrelated phenological, morphological, and physiological traits. It is these small but coordinated differences in the plastic responses of many traits that appear to differentiate cocklebur from natural vs. weedy urban habitats.