Competition under high and low nutrient levels among three grassland species occupying different positions in a successional sequence

To clarify the role of seasonal change, competitive response and nutrient availability in the competitive asymmetry of grassland species a competition experiment was conducted on Holcus lanatus , Anthoxanthum odoratum and Festuca ovina , which represent a successional sequence of decreasing nutrient availability. Seven harvests were taken over two growing seasons. At each harvest the dry weight of plant parts, dead leaves, leaf area and plant height were measured. Three key traits that determine the successional status of the species were studied: specific leaf area, specific shoot height, and dead leaf fraction.
The response of these traits to competition appeared to be limited and insufficient to change the competitive relations in the experiment. However, all three traits showed marked seasonal changes which resulted in superior growth and survival in winter of the species adapted to nutrient-poor environments. The findings support the theory that competitive asymmetry increases at higher nutrient levels. It is postulated that the directionality of light makes it possible for the dominant species to monopolize this resource more easily than nutrients.     

Growth,biomass allocation and plant nitrogen concentration in Chilean temperate rainforest tree seedlings: effects of nutrient availability

Seedlings of nine southern Chilean trees were grown at three nutrient supply rates, to examine the roles of growth rate, biomass distribution and nutrient use traits in determining species natural distributions on resource gradients. Relative growth rate (RGR) showed no overall relationship with species site requirements, although RGR of fertile-site species tended to be more responsive to nutrient supply. In the low-nutrient treatment, infertility-tolerant Fitzroya cupressoides showed a higher RGR rank than a fertility-demanding species (Laurelia philippiana) which outgrew it substantially at the highest supply rate. This reversal of RGR ranks was associated with divergent nutrient use responses: at high nutrient supply both spp. had similar plant nitrogen concentrations (PNC), whereas at the low supply rate Fitzroya’s production of biomass per unit of assimilated N was twice that of Laurelia’s. However, this pattern does not appear to serve as a general explanation of the respective distributions of the study species, as RGR ranks of most species were unaltered by nutrient supply. At low nutrient availability, no clear differences in shoot:root ratio (SRR) were apparent between poor-site and fertile-site species. However, at high nutrient availability, SRR was markedly higher in the latter, resulting from differences in biomass allocation to stems (not leaves). Leaf area ratios (LAR) were higher in fertile-site species than in those tolerant of low fertility, because of differences in specific leaf area rather than leaf weight ratio. Very high LAR at high nutrient supply was characteristic of most shade-tolerant angiosperms, but not of shade-tolerant conifers. Although PNC showed no overall differences between poor- and fertile-site species, sensitivity of PNC to external supply rate was greatest in two infertility-tolerant conifers. In contrast, the angiosperm Weinmannia trichosperma, although tolerant of low fertility, responded to increased nutrient supply with greatly increased RGR and little change in PNC. Results show little trait convergence between conifers and angiosperms in adaptation both to shade and to infertile soils; i.e. fitness of different taxa in a given environment may hinge on different trait combinations. Received: 12 September 1995 /Accepted: 14 June 1996     

Root and leaf attributes accounting for the performance of fast- and slow-growing grasses at different nutrient supply

Despite their difference in potential growth rate, the slow-growing Brachypodium pinnatum and the fast-growing Dactylis glomerata co-occur in many nutrient-poor calcareous grasslands. They are known to respond differently to increasing levels of N and P. An experiment was designed to measure which characteristics are affected by nutrient supply and contribute to the ecological performance of these species. Nutrient acquisition and root and shoot traits of these grasses were studied in a garden experiment with nine nutrient treatments in a factorial design of 3 N and 3 P levels each. D. glomerata was superior to B. pinnatum in nutrient acquisition and growth in all treatments. B. pinnatum was especially poor in P acquisition. Both species responded to increasing N supply and to a lesser extent to increasing P supply by decreasing their root length and increasing their leaf area per total plant weight. D. glomerata showed a higher plasticity. In most treatments, the root length ratio (RLR) and the leaf area ratio (LAR) were higher for D. glomerata. A factorization of these parameters into components expressing biomass allocation, form (root fineness or leaf thickness) and density (dry matter content) shows that the low density of the biomass of D. glomerata was the main cause for the higher RLR and LAR. The biomass allocation to the roots showed a considerable plasticity but did not differ between the species. B. pinnatum had the highest leaf weight ratio. Root fineness was highly plastic in D. glomerata, the difference with B. pinnatum being mainly due to the thick roots of D. glomerata at high nutrient supply. The leaf area/leaf fresh weight ratio did not show any plasticity and was slightly higher for B. pinnatum. It is concluded, that the low density of the biomass of D. glomerata is the pivotal trait responsible for its faster growth at all nutrient levels. It enables simultaneously a good nutrient acquisition capacity by the roots as well as a superior carbon acquisition by the leaves. The high biomass density of B. pinnatum will then result in a lower nutrient requirement due to a slower turnover, which in the long term is advantageous under nutrient-poor conditions.     

Allocation,within and between organs,and the dynamics of root length changes in two birch species

Spatial and temporal dynamics of biomass allocation within and between organs were investigated in seedlings of two birch species of contrasting successional status. Seedlings of Betula alleghaniensis Britt (yellow birch) and B. populifolia Marsh (gray birch) were grown for 6 weeks at two nutrient levels in rectangular plexiglass containers to allow non-destructive estimates of root growth, production and loss. Leaf area and production were simultaneously monitored. Yellow birch responded more to nutrient level than gray birch in terms of total biomass, shoot biomass, leaf area and root length. Yellow birch also flexibly altered within-organ allocation (specific leaf area, specific root length and specific soil amount). In contrast, gray birch altered between-organ allocation patterns (root length:leaf area and soil amount:leaf area ratios) more than yellow birch in response to nutrient level. Yellow birch showed greater overall root density changes within a very compact root system, while gray birch showed localized root density changes as concentric bands of new root production spread through the soil. Species differ critically in their responses of standing root length and root production and loss rates to nutrient supply. Early successional species such as gray birch are hypothesized to exhibit higher plasticity in varied environments than later successional species such as yellow birch. Our results suggest that different patterns of allocation, within and between plant organs, do not necessarily follow the same trajectories. To characterize thoroughly the nature of functional flexibility through ontogeny, within- and between-organ patterns of allocation must be accounted for.     

Traits of an invasive grass conferring an early growth advantage over native grasses

Aims Invasive species often have higher relative growth rates (RGR) than their native counterparts. Nutrient use efficiency, total leaf area and specific leaf area (SLA) are traits that may confer RGR differences between natives and invasives, but trait differences are less prominent when the invasive species belongs to the same plant functional type as the dominant native species. Here, we test if traits displayed soon after germination confer an early size advantage. Specifically, we predicted that invasive species seedlings grow faster than the natives because they lack trade-offs that more strongly constrain the growth of native species.Methods We quantified plant morphological and physiological traits and RGR during early seedling growth at high and low nutrient levels in three dominant perennial native C₄ grasses: Panicum virgatum L. (switchgrass), Schizachyrium scoparium (Michx.) Nash (little bluestem) and Andropogon gerardii Vitman (big bluestem); and a perennial C₄ exotic invasive grass, Sorghum halepense (L.) Pers. (Johnsongrass).Important findings After 2 weeks of growth, Johnsongrass seedlings had greater biomass, SLA and photosynthetic nitrogen use efficiency, but lower leaf N concentrations (% leaf N) and root:shoot ratio than natives. As growth continued, Johnsongrass more quickly produced larger and thicker leaves than the natives, which dampened the growth advantage past the first 2 to 3 weeks of growth. Investment in carbon gain appears to be the best explanation for the early growth advantage of Johnsongrass. In natives, growth was constrained by an apparent trade-off between allocation to root biomass, which reduced SLA, and production of leaves with high N content, which increased carbon gain. In Johnsongrass, root:shoot ratio did not interact with other traits, and % leaf N was decoupled from RGR as a result of a trade-off between the positive indirect association of % leaf N with RGR and the negative direct association of % leaf N with RGR.     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

Biomass estimation models and allocation patterns of 14 shrub species in Mountain Luya,Shanxi, China

Aims Shrub species have evolved specific strategies to regulate biomass allocation among various organs or between above- and belowground biomass and shrub biomass model is an important approach to estimate biomass allocation among different shrub species. This study was designed to establish the optimal estimation models for each organ (leaf, stem, and root), aboveground and total biomass of 14 common shrub species in Mountain Luya, Shanxi Province, China. Furthermore, we explored biomass allocation characteristics of these shrub species by using the index of leaf biomass fraction (leaf to total biomass), stem biomass fraction (stem to total biomass), root biomass fraction (root to total biomass), and root to shoot mass ratio (R/S) (belowground to aboveground biomass).
Methods We used plant height, basal diameter, canopy diameter and their combination as variables to establish the optimal biomass estimation models for each shrub species. In addition, we used the ratios of leaf, stem, root to total biomass, and belowground to aboveground biomass to explore the difference of biomass allocation patterns of 14 shrub species.
Important findings Most of biomass estimation models could be well expressed by the exponential and linear functions. Biomass for shorter shrub species with more stems could be better estimated by canopy area; biomass for taller shrub species with less stems could be better estimated by the sum of the square of total base diameter multiply stem height; and biomass for the rest shrub species could be better estimated by canopy volume. The averaged value for these shrub species was 0.61, 0.17, 0.48, and 0.35 for R/S, leaf biomass fraction, stem biomass fraction, and root biomass fraction, respectively. Except for leaf biomass fraction, R/S, stem biomass fraction, and root biomass fraction for shrubs with thorn was significantly greater than that for shrubs without thorn.     

三种高寒植物幼苗生物量分配及性状特征对光照和养分的响应

为了解高寒植物幼苗对生境资源异质性的适应策略,以高寒草甸中常见的3种草本植物大耳叶风毛菊(Saussurea macrota)、甘西鼠尾草(Salvia przewalskii)和千里光(Senecio scandens)为材料,比较研究了这3种植物幼苗对不同光照和养分资源的响应。结果表明:光照和养分异质性显著影响了3种植物幼苗的性状特征和生物量分配,并存在一定的交互影响。随着光照的降低,3个物种的幼苗的生物量和根分配呈现降低趋势,但是其株高、比叶面积、叶分配、茎分配却逐渐升高。在低养分条件下,3个物种幼苗的总生物量、株高、比叶面积和叶分配均降低,而根分配均却显著增加。对于光照和养分资源异质性而言,光照异质性对高寒植物生物量分配和性状特征的改变具有更大的影响。喜阴物种大耳叶风毛菊和喜光物种甘西鼠尾草比中性生境物种千里光表现出了较大的性状特征和生物量分配的可塑性指数。     

Interspecific Variation in RGR and the Underlying Traits among 24 Grass Species Grown in Full Daylight

Abstract: A growth analysis was conducted with 24 central European grass species in full daylight to test whether traits underlying interspecific variation in relative growth rate (RGR) are the same in full daylight as they are at lower light, and whether this depends on the ecological characteristics of the studied species, i.e., their requirements with respect to nutrient and light availability.
In contrast to studies with herbaceous species at lower light, net assimilation rate (NAR) contributed more than leaf area ratio (LAR) or specific leaf area (SLA) to interspecific variation in RGR. This was associated with a larger interspecific variation in NAR than found in experiments with lower light. Without the two most shade-tolerant species, however, the contribution of LAR and its components to interspecific variation in RGR was similar or even higher than that of NAR.
Leaf dry matter content correlated negatively with RGR and was the only component of LAR contributing in a similar manner to variation in LAR and RGR. There was a positive correlation between NAR and biomass allocation to roots, which may be a result of nutrient-limited growth. RGR correlated negatively with biomass allocation to leaves. Leaf thickness did not correlate with RGR, as the positive effect of thin leaves was counterbalanced by their lower NAR.
Low inherent RGR was associated with species from nutrient-poor or shady habitats. Different components constrained growth for these two groups of species, those from nutrient-poor habitats having high leaf dry matter content, while those from shady habitats had thin leaves with low NAR.     

Intra-specific variation in relative growth rate: impact on competitive ability and performance of Lychnis flos-cuculi in habitats differing in soil fertility

Plant species from unproductive or adverse habitats are often characterized by a low potential relative growth rate (RGR). Although it is generally assumed that this is the result of selection for specific trait combinations that are associated with a low rate of net biomass accumulation, few studies have directly investigated the selective (dis-)advantage of specific growth parameters under a set of different environmental conditions. Aim of the present study was to quantify the impact of inherent differences in growth parameters among phenotypes of a single plant species, Lychnis flos-cuculi, on their performance under different soil nutrient conditions. Growth analysis revealed significant variation in RGR among progeny families from a diallel cross between eight genotypes originating from a single population. Differences in RGR were due to variation in both leaf area ratio (LAR) and in net assimilation rate (NAR). A genetic trade-off was observed between these two components of growth, i.e. progeny families with high investment in leaf area had a lower rate of net biomass accumulation per unit leaf area. The degree of plasticity in RGR to nutrient conditions did not differ among progeny families. Inherent differences in growth parameters among progeny families had a significant impact on their yield in competition with Anthoxanthum odoratum and Taraxacum hollandicum. In nutrient-rich conditions, progeny families with an inherently high leaf weight ratio (LWR) achieved higher yield in competition, but variation in this trait could not explain differences in competitive yield under nutrient-poor conditions. Inherent differences in growth parameters among progeny families were poorly correlated with differences in survival and average rosette biomass (a good predictor of fecundity) among these progeny families sown in four field sites along a natural gradient of soil fertility. In the more productive sites none of the growth parameters was significantly correlated with rosette biomass, but in the least productive site progeny families with an inherently high specific leaf area (SLA) tended to produce smaller rosettes than low-SLA families. These results are consistent with the view that a selective advantage may accrue from either high or low values of individual RGR components, depending on habitat conditions, and that the selective advantage of low trait values in nutrient-poor environments may results in indirect selection for low RGR in these habitats.     

Phenotypic plasticity of <Emphasis Type="Italic">Chenopodium murale</Emphasis> across contrasting habitat conditions in peri-urban areas in Indian dry tropics: Is it indicative of its invasiveness?

Phenotypic plasticity is an important plant trait associated with invasiveness of alien plants that reflects its ability to occupy a wide range of environments. We investigated the phenotypic response of Chenopodium murale to resource variability and ontogeny. Its plant-level and leaf-level traits were studied at high-resource (HR) and low-resource (LR) sites in peri-urban areas in Indian dry tropics. Plants at LR had significantly higher root length, root/shoot biomass ratio, stem mass and root mass fractions. Plants at HR had higher shoot length, basal diameter, leaf mass fraction and leaf area ratio. Leaf-level traits like leaf area and chlorophyll a were also higher here. Mean plasticity indices for plant- and leaf-level traits were higher at HR. With increasing total plant biomass, there was significant increase in the biomass of leaf, stem, root, and reproductive parts, and root and shoot lengths, whereas root/shoot length ratio, their biomass ratio, and leaf and root mass fractions declined significantly. Allocation to roots and leaves significantly decreased with increasing plant size at both sites. But, at any size, allocation to roots was greater at LR, indicative of optimization of capture of soil nutrients, whereas leaf allocation was higher at HR. Consistently increasing stem allocation equaled leaf allocation at comparatively higher shoot lengths at HR. Reproductive biomass comprised 10–12% of the plant’s total biomass. In conclusion, the success of alien weed C. murale across environmentally diverse habitat conditions in Indian dry tropics can be attributed to its high phenotypic plasticity, resource utilization capability in low-resource habitats and higher reproductive potential. These characteristics suggest that it will continue to be an aggressive invader.     

Allometry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan

Plant allometry that is related to plant architecture and biomass allocation strongly influences a plants ability to grow in shaded forest understory. Some allometric traits can change with plant size. The present study compared crown and trunk allometries, root/shoot biomass allometry, and root architecture among understory saplings (0.25--5m height, except for two trees > 5 < 7 m) of seven deciduous dicotyledonous species in central Japan. Associations of the crown and trunk allometries with several plant morphological attributes were analyzed. Branch morphology (plagiotropyvs orthotropy) and life size were correlated with sapling crown and trunk allometries. Both large leaves and orthotropic branches were associated with a narrow small crown and slender trunk. The root/shoot ratio decreased rapidly with increasing plant height for saplings shorter than about 1.5 m. Less shade-tolerant species tended to have smaller root/shoot ratios for saplings taller than 1.5 m. With an increase in plant height, the branch/trunk biomass ratio decreased for saplings with plagiotropic branches but increased for saplings with orthotropic branches. Four subcanopy species (Acer distylum, Carpinus cordata, Fraxinus lanuginosa and Acanthopanax sciadophylloides) had superficial root systems; a common understory species (Sapium japonica) had a deep tap root system; and a canopy species (Magnolia obovata) and a subcanopy species (Acer tenuifolium) had heart root systems of intermediate depth. The root depth was not related to shade tolerance. Among species of the same height, the difference in fine root length can be 30-fold.     

Growth,biomass allocation,morphology, and photosynthesis of invasive <Emphasis Type="Italic">Eupatorium</Emphasis><Emphasis Type="Italic">adenophorum</Emphasis> and its native congeners grown at four irradiances

Eupatorium adenophorum is one of the more noxious invasive plants worldwide. However, the mechanisms underlying its invasiveness are still not well elucidated. In this study, we compared the invader with its two native congeners (E. heterophyllum and E. japonicum) at four irradiances in terms of growth, biomass allocation, morphology, and photosynthesis. The higher light-saturated photosynthetic rate (P _max) and total leaf area of the invader may contribute to its higher relative growth rate (RGR) and total biomass compared with its native congeners. Total biomass and RGR increased significantly with the increase of P _max and total leaf area. The higher support organ mass fraction and the lower root mass fraction of the invader may also contribute to its higher RGR and biomass through increasing carbon assimilation and reducing respiratory carbon loss, respectively. The higher growth rate of the invader increased its total leaf area, ramet number, and crown area. These traits may help the invader to form dense monoculture, outshading native plant species. However, consistently higher leaf area ratio, specific leaf area, and leaf mass fraction were not found across irradiances for the invader compared with its native congeners. Higher plasticity in response to irradiance was also not found for the invader. The invader retained advantages over the natives across irradiances, while its performance decreased with lower irradiance. The results indicate that the invader may be one of the few super invaders. Reducing irradiance may inhibit its invasions.     

Explaining patterns of primary production from individual level traits

Question: Do species traits explain differences in productivity in grazed and ungrazed plots? Location: Río de la Plata grasslands, Uruguay (31°54′S, 58°15′W). Methods: In a greenhouse experiment, we measured the relative growth rate (RGR) of grasses with contrasting responses to grazing (increasers and decreasers). We performed six harvests at weekly intervals in order to calculate the RGR and assess 12 plant traits. We compared the RGR between increaser and decreaser species after 2 and 5 weeks using t‐tests. Linear and potential regression models were fitted to time versus natural logarithm of total dry biomass relationships. The RGR temporal trajectories of increaser and decreaser species were obtained from the derivatives of the best‐fit functions. Principal component analysis (PCA) was used to sort species according to their traits. Results: The RGR of decreaser grasses was higher than that of increasers at the second week, while at the fifth week the opposite was observed. The RGR of decreasers dropped through time, while the RGR of increaser species was constant. The PCA separated increaser from decreaser species. The attributes related to increaser species were: high specific leaf area, tillering rate, green leaf rate, total leaf number, root weight ratio and leaf weight ratio; while those associated with decreaser species were: high dead biomass, senescence rate, reproductive biomass, leaf elongation rate and total biomass. Conclusions: Traits possessed by decreasers reduce light availability and increase the reproductive investment, explaining the drop in RGR. Specific differences in RGR seem to scale up to the ecosystem level and would explain the pattern in aboveground net primary production observed in the field under contrasting grazing regimes.     

Leaf area ratio and net assimilation rate of 24 wild species differing in relative growth rate

Summary Which factors cause fast-growing plant species to achieve a higher relative growth rate than slow-growing ones? To answer this question 24 wild species were grown from seed in a growth chamber under conditions of optimal nutrient supply and a growth analysis was carried out. Mean relative growth rate, corrected for possible ontogenetic drift, ranged from 113 to 356 mg g^–1 day^–1. Net assimilation rate, the increase in plant dry weight per unit leaf area and unit time, varied two-fold between species but no correlation with relative growth rate was found. The correlation between leaf area ratio, the ratio between total leaf area and total plant weight, and relative growth rate was very high. This positive correlation was mainly due to the specific leaf area, the ratio between leaf area and leaf weight, and to a lesser extent caused by the leaf weight ratio, the fraction of plant biomass allocated to the leaves. Differences in relative growth rate under conditions of optimum nutrient supply were correlated with the soil fertility in the natural habitat of these species. It is postulated that natural selection in a nutrient-rich environment has favoured species with a high specific leaf area and a high leaf weight ratio, and consequently a high leaf area ratio, whereas selection in nutrient-poor habitats has led to species with an inherently low specific leaf area and a higher fraction of root mass, and thus a low leaf area ratio.     

南亚热带森林24种乔木的种子萌发和幼苗生长

以膨胀珍珠岩为基质,在光和暗的条件下,对２４种南亚热带森林乔木的种子萌发和幼苗生长进行了研究。种子的形态和重量与种的演替阶段有关,种子的重量也与不同的种有关。不同种的萌发率差异较大,种子较大的种,萌发率较高。肉质果实的种子,开始萌发的时间较长。光和暗条件对萌发率和萌发速度无明显影响。幼苗高度和种子重量呈正相关,这一相关在光条件下比暗条件下更明显。暗条件明显地引起群落演替早期的树种的茎徒长,限制根的生长,而对演替后期种无显著的影响。幼苗地上部分和根系生物量的分配与种的演替阶段有关,也与不同的科有关。幼苗地上部分和根系生物量的分配也受光暗条件的影响,而不受种子重量的影响。根和叶的生物量分配和种在其群落演替阶段有关。幼苗的生物量与种子重量有显著的正相关。幼苗的相对生长率相差较大,在没有营养供应的生长基质中,以演替中间阶段的种的相对生长率较大。幼苗从种子的物质利用效率与种子重量呈负相关,而与种的演替阶段无明显的相关。     

Influence of light and nutrient conditions on seedling growth of native and invasive trees in the Seychelles

Several recent studies have shown that plant invasions can occur in resource-poor and relatively undisturbed habitats. It is, therefore, important to investigate whether and how life-history traits of species invasive in such habitats differ from those of species that are only invasive in disturbed and resource rich habitats. We compared the growth of seedlings of native and invasive tree species from nutrient-poor secondary forests in the tropical Seychelles. We hypothesised that the relative performance of the two groups would change predictably along resource gradients, with native species performing better at low levels of resource availability and invasive species performing better at higher levels. To test this hypothesis, we performed a common garden experiment using seedlings of six invasive and seven native tree species grown under three levels of light (65, 11 and 3.5% of ambient light) and two of nutrients (low and high). Due to large variation among species, differences in growth rates (RGR) were not significant among seedlings of the native and the invasive species. However, seedlings of the invasive species showed higher specific leaf areas (SLA) and higher leaf nutrient contents than seedlings of the native species. They also exhibited greater plasticity in biomass and nutrient allocation (i.e., greater plasticity in LAR, RSR and leaf nutrient contents) in response to varying resource availability. However, differences between the mean values of these parameters were generally small compared with variation within groups. We conclude that successful invaders on nutrient-poor soils in the Seychelles are either stress-tolerant, possessing growth traits similar to those of the native species, or fast-growing but adapted to nutrient-poor soils. In contrast, the more typical, fast-growing alien species with no particular adaptations to nutrient-poor soils seem to be restricted to relative nutrient-rich sites in the lowlands. The finding—that some introduced species thrive in resource-poor habitats—suggests that undisturbed habitats with low resource availability may be less resistant to plant invasions than was previously supposed.     

Root architecture and allocation patterns of eight native tropical species with different successional status used in open-grown mixed plantations in Panama

We investigated biomass allocation and root architecture of eight tropical species with different successional status, as classified from the literature, along a size gradient up to 5 m. We focused on belowground development, which has received less attention than aboveground traits. A discriminant analysis based upon a combination of allocational and architectural traits clearly distinguished functional types and classified species according to successional status at a 100% success rate. For a given plant diameter, the pioneer species presented similar root biomass compared to the non-pioneer ones but higher cumulative root length and a higher number of root apices. A detailed study on the root system of a sub-sample of three species showed that the most late-successional species (Tabebuia rosea) had longer root internodes and a higher proportion of root biomass allocated to the taproot compared to the other two species (Hura crepitans and Luehea seemannii). Most pioneer species showed a higher leaf area ratio due to a higher specific leaf area (SLA). We conclude that the functional differences between pioneer and non-pioneer tree species found in natural forests were maintained in open-grown plantation conditions.     

Biomass allocation in old-field annual species grown in elevated CO2 environments: no evidence for optimal partitioning

Increased atmospheric carbon dioxide supply is predicted to alter plant growth and biomass allocation patterns. It is not clear whether changes in biomass allocation reflect optimal partitioning or whether they are a direct effect of increased growth rates. Plasticity in growth and biomass allocation patterns was investigated at two concentrations of CO₂ ([CO₂]) and at limiting and nonlimiting nutrient levels for four fast‐ growing old‐field annual species. Abutilon theophrasti, Amaranthus retroflexus, Chenopodium album, and Polygonum pensylvanicum were grown from seed in controlled growth chamber conditions at current (350 μmol mol^?1, ambient) and future‐ predicted (700 μmol mol^?1, elevated) CO₂ levels. Frequent harvests were used to determine growth and biomass allocation responses of these plants throughout vegetative development. Under nonlimiting nutrient conditions, whole plant growth was increased greatly under elevated [CO₂] for three C3 species and moderately increased for a C4 species (Amaranthus). No significant increases in whole plant growth were observed under limiting nutrient conditions. Plants grown in elevated [CO₂] had lower or unchanged root:shoot ratios, contrary to what would be expected by optimal partitioning theory. These differences disappeared when allometric plots of the same data were analysed, indicating that CO₂‐induced differences in root:shoot allocation were a consequence of accelerated growth and development rates. Allocation to leaf area was unaffected by atmospheric [CO₂] for these species. The general lack of biomass allocation responses to [CO₂] availability is in stark contrast with known responses of these species to light and nutrient gradients. We conclude that biomass allocation responses to elevated atmospheric [CO₂] are not consistent with optimal partitioning predictions.     

The effects of clipping and soil moisture on leaf and root morphology and root respiration in two temperate and two tropical grasses

Numerous studies have explored the effect of environmental conditions on a number of plant physiological and structural traits, such as photosynthetic rate, shoot versus root biomass allocation, and leaf and root morphology. In contrast, there have been a few investigations of how those conditions may influence root respiration, even though this flux can represent a major component of carbon (C) pathway in plants. In this study, we examined the response of mass-specific root respiration (μmol CO₂ g⁻¹ s⁻¹), shoot and root biomass, and leaf photosynthesis to clipping and variable soil moisture in two C₃ (Festuca idahoensis Elmer., Poa pratensis L.) and two C₄ (Andropogon greenwayi Napper, and Sporobolus kentrophyllus K. Schum.) grass species. The C₃ and C₄ grasses were collected in Yellowstone National Park, USA and the Serengeti ecosystem, Africa, respectively, where they evolved under temporally variable soil moisture conditions and were exposed to frequent, often intense grazing. We also measured the influence of clipping and soil moisture on specific leaf area (SLA), a trait associated with moisture conservation, and specific root length (SRL), a trait associated with efficiency per unit mass of soil resource uptake. Clipping did not influence any plant trait, with the exception that it reduced the root to shoot ratio (R:S) and increased SRL in P. pratensis. In contrast to the null effect of clipping on specific root respiration, reduced soil moisture lowered specific root respiration in all four species. In addition, species differed in how leaf and root structural traits responded to lower available soil moisture. P. pratensis and A. greenwayi increased SLA, by 23% and 33%, respectively, and did not alter SRL. Conversely, S. kentrophyllus increased SRL by 42% and did not alter SLA. F. idahoensis responded to lower available soil moisture by increasing both SLA and SRL by 38% and 33%, respectively. These responses were species-specific strategies that did not coincide with photosynthetic pathway (C₃/C₄) or growth form. Thus, mass-specific root respiration responded uniformly among these four grass species to clipping (no effect) and increased soil moisture stress (decline), whereas the responses of other traits (i.e., R:S ratio, SLA, SRL) to the treatments, especially moisture availability, were species-specific. Consequently, the effects of either clipping or variation in soil moisture on the C budget of these four different grasses species were driven primarily by the plasticity of R:S ratios and the structural leaf and root traits of individual species, rather than variation in the response of mass-specific root respiration.