Relative growth rate and biomass allocation in ten woody species with different leaf longevity using phylogenetic independent contrasts (PICs)

In this study, we compare the relative growth rate (RGR) and biomass allocation of 10 woody species (5 deciduous and 5 evergreen) from the Mediterranean region using phylogenetic independent contrasts (PICs) to test if these two functional groups differ in these traits. In general, the results were similar when using PICs or without taking into account phylogenetic relations. Deciduous species had a higher RGR than evergreen species, due to the higher net assimilation rate (NAR). Deciduous species had a higher specific leaf area (SLA) but a lower leaf mass ratio (LMR), resulting in a similar LAR for deciduous and evergreen species (LAR = SLA x LMR). In some cases, the use of PICs revealed patterns that would not have appeared if phylogeny had been overlooked. For example, there was no significant correlation between RGR and final dry mass (after 4 months of growth) but PICs revealed that there was a positive relation between these two variables in all deciduous-evergreen pairs. In general, RGR decreased with time and this temporal variation was due primarily to NAR variations (r = 0.79, p < 0.01), and also to variations in LAR (r = 0.69, p < 0.05). Considering the phylogeny, the only variable constantly different for all deciduous-evergreen pairs was SLA. This result, and the fact that SLA was the best correlated variable with RGR (r = 0.81, p < 0.01), reinforce the value of SLA as a variable closely associated to growth and to the functional groups (deciduous vs. evergreen).     

Net Assimilation Rate Determines the Growth Rates of 14 Species of Subtropical Forest Trees

Growth rates are of fundamental importance for plants, as individual size affects myriad ecological processes. We determined the factors that generate variation in RGR among 14 species of trees and shrubs that are abundant in subtropical Chinese forests. We grew seedlings for two years at four light levels in a shade-house experiment. We monitored the growth of every juvenile plant every two weeks. After one and two years, we destructively harvested individuals and measured their functional traits and gas-exchange rates. After calculating individual biomass trajectories, we estimated relative growth rates using nonlinear growth functions. We decomposed the variance in log(RGR) to evaluate the relationships of RGR with its components: specific leaf area (SLA), net assimilation rate (NAR) and leaf mass ratio (LMR). We found that variation in NAR was the primary determinant of variation in RGR at all light levels, whereas SLA and LMR made smaller contributions. Furthermore, NAR was strongly and positively associated with area-based photosynthetic rate and leaf nitrogen content. Photosynthetic rate and leaf nitrogen concentration can, therefore, be good predictors of growth in woody species.     

Contribution of root growth responses to leaf traits and relative growth rate of Populus alba under different water-table conditions

Water-table depth variations alter root growth response and may affect whole-plant growth in arid and semi-arid regions. We examined how root biomass allocation and root morphological traits affect the leaf physiological and morphological traits and whole-plant growth of Populus alba growing under different water tables. We exposed 1-year-old P. alba cuttings to contrasting soil–water conditions via water table changes in a greenhouse for 90 days. We examined relationships among net assimilation rate (NAR) and other growth components obtained from our published data for trees harvested every 30 days. Strongly negative correlations were found between RMR and root morphological traits. Root mass ratio had a strong negative relationship with LMR, and proportion of fine-root biomass per total root biomass was positively correlated with SLA and NAR. Both NAR and leaf area ratio were important determinants of variation in relative growth rate (RGR). Leaf mass ratio (LMR) and specific leaf area (SLA) were positively correlated with RGR; the correlation was stronger in the case of LMR. Along a water-table gradient, negative relationships between root growth responses are likely to indirectly influence RGR through changes in NAR, LMR, and SLA.     

Relative growth rate in phylogenetically related deciduous and evergreen woody species

Relative growth rate (RGR) and other growth parameters were studied in eight pairs of closely related deciduous and evergreen species (within the same genus or family). The main objective of this study was to test the association between leaf turnover rate and RGR, specific leaf area (SLA, leaf area/leaf dry weight) and other growth variables. Plants were grown for 6 months in a greenhouse under favourable water and nutrient conditions. Variation in RGR among the 16 woody species was due mainly to differences in morphological parameters such as leaf area ratio (LAR, whole plant area/whole plant dry weight) and SLA). However, temporal variation in RGR within species was due mainly to variation in net assimilation rate. When phylogeny was not taken into account, analyses showed that deciduous species grew faster than evergreens. In contrast, when phylogeny was taken into account, the data analysis showed that a faster RGR is not consistently associated with the deciduous habit (in five pairs it was, but in the other three it was not). The faster growth of the deciduous trees (in the five positive contrasts) could be explained by their higher LAR and higher SLA relative to evergreens. The lack of differences in RGR between deciduous and evergreens (in three pairs) was due to the higher leaf mass ratio (LMR, leaf dry biomass/total dry biomass) for the evergreens, which offset the higher SLA of the deciduous species, resulting in a similar LAR in both functional groups (LAR=LMR2SLA). Deciduous species had consistently higher SLA than evergreens. We suggest that SLA, more than RGR, could be an important parameter in determining adaptive advantages of deciduous and evergreen species.     

Variation in relative growth rate of 20 Aegilops species (Poaceae) in the field: The importance of net assimilation rate or specific leaf area depends on the time scale

Field experiments reporting the relative growth rate (RGR) patterns in plants are scarce. In this study, 22 herbaceous species (20 Aegilops species, Amblyopyrum muticum and Triticum aestivum) were grown under field conditions to assess their RGR, and to find out if the differences in RGR amongst species were explained by morphological or physiological traits. Plants were cultivated during two months, and five harvests (every 13–19 days) were carried out. Factors explaining between-species differences in RGR varied, depending on whether short (13–19 days) or longer periods (62 days) were considered. RGR for short periods (4 growth periods of 13–19 days each) showed a positive correlation with net assimilation rate (NAR), but there was no significant correlation with leaf area ratio (LAR) (with the exception of the first growth period). In contrast, when growth was investigated over two months, RGR was positively correlated with morphological traits (LAR, and specific leaf area, SLA), but not with physiological traits (NAR). A possible explanation for these contrasting results is that during short growth periods, NAR exhibited strong variations possibly caused by the variable field conditions, and, consequently NAR mainly determined RGR. In contrast, during a longer growth period (62 days) the importance of NAR was not apparent (there was no significant correlation between RGR and NAR), while allocation traits, such as LAR and SLA, became most relevant.     

Plant functional traits shape growth rate for xerophytic shrubs

• Trade-offs exist for xerophytic shrubs between functional traits, involving in water loss and assimilate accumulation, can contribute to its survival and growth rate regulation in arid environments. However, growth analysis based on plant functional traits has been focused on the study of herbs and woody species. It is still unclear how the functional traits of xerophytic shrubs regulate their growth rate.
• In this study, we selectedeight xerophytic shrubs as samples to analyze the regulation process of the functional traits of shrubs on growth rate. Plants were cultivated for three years, and three harvests (every one year) were carried out. Factors explaining between-species differences in relative growth rate (RGR) varied, depending on whether different ages were considered.
• The results showed that RGR was positively correlated with net assimilation rate, but there was a significant negative correlation with leaf area ration (LAR), specific leaf area (SLA), and leaf biomass ratio in the age 1. However, in the age 2, RGR showed a significant positive correlation with the morphological traits (i.e., leaf area ration and specific leaf area), but not with physiological traits (i.e., net assimilation rate) and leaf biomass allocation.
• Our results suggested that the fluctuation of environmental factors affects the regulation path of the plant functional traits on RGR of xerophytic shrubs. However, the analysis of causality model showed that no matter in which age, net assimilation rate and leaf area ration principally drive the variation in RGR among xerophytic shrubs.

     

Interspecific correlates of plasticity in relative growth rate following a decrease in nitrogen availability

Background and Aims

Nitrogen availability varies greatly over short time scales. This requires that a well-adapted plant modify its phenotype by an appropriate amount and at a certain speed in order to maximize growth and fitness. To determine how plastic ontogenetic changes in each trait interact and whether or not these changes are likely to maximize growth, ontogenetic changes in relative growth rate (RGR), net assimilation rate (NAR), specific leaf area (SLA) and root weight ratio (RWR), before and after a decrease in nitrogen supply, were studied in 14 herbaceous species.

Methods

Forty-four plants of each species were grown in hydroponic culture under controlled conditions in a control treatment where the supply of nitrogen remained constant at 1 mm, and in a stress treatment where the nitrogen supply was abruptly decreased from 1 to 0·01 mm during the growth period.

Key Results and Conclusions

In the treatment series, and in comparison with the control, NAR and RGR decreased, RWR increased, and SLA did not change except for the timing of ontogenetic change. Species having greater increases in the maximum rate of change in RWR also had smaller reductions in RGR; plasticity in RWR is therefore adaptive. In contrast, species which showed a greater decrease in NAR showed stronger reductions in RGR; plasticity in NAR is therefore not adaptive. Plasticity in RGR was not related to plasticity in SLA. There were no significant relationships among the plasticities in NAR, RWR or SLA. Potentially fast-growing species experienced larger reductions in RGR following the nitrogen reduction. These results suggest that competitive responses to interspecific competition for nitrogen might be positively correlated with the plasticity in the maximum rate of change in RWR in response to a reduction in nitrogen supply.     

Understanding seedling growth relationships through specific leaf area and leaf nitrogen concentration: generalisations across growth forms and growth irradiance

Seedling relative growth rate (RGR) achieved under favourable growth conditions can be thought of as a useful bioassay of the potential ability of species to take advantage of favourable growth opportunities; that is, of a species' growth strategy. The consistency of relationships between RGR and its component attributes leaf nitrogen productivity (LNP), leaf N per area (LNCa), specific leaf area (SLA) and leaf mass ratio (LMR) was assessed across 12 datasets comprising three growth forms (grasses, herbaceous dicots and woody plants; 250 species in total). These relationships were characterised in terms of scaling slopes (regressions on log-log axes, the slopes giving the proportional relationship between the variables). Mathematically, the expected scaling slope between RGR and each component is 1.0, giving an appropriate null hypothesis to test against (whereas the widely used null hypothesis of zero correlation is in fact inappropriate for this situation). Deviations below 1:1 scaling slopes indicate negative covariance between the components. Consequently, the correlation structure between the components of RGR should also be investigated. Biologically, RGR should scale 1:1 with SLA at a given LNCa and somewhat more weakly with LNCa at a given SLA. SLA and LNCa should themselves scale with a slope of between 0 and -1, with the actual slope indicating the extent to which between-species variation in SLA dilutes leaf N on an area basis versus the ability of species to maintain LNCa at a given growth irradiance. On average, across the 12 datasets RGR scaled close-to-proportionally with SLA, and 1:1 with SLA at a given LNCa. RGR scaled with LNCa with null or negative slopes, since SLA and LNCa scaled negatively (with slopes generally shallower than -1); however, RGR scaled positively (but less than proportionally) with LNCa at a given SLA. For these key relationships there were no qualitatively different conclusions with respect to the growth form under consideration or the growth irradiance at which the seedlings were grown. RGR also scaled close-to-proportionally with LNP, while LNP and LNCa were negatively associated. These relationships involving LNP are difficult to interpret since it can be shown that they are, at least potentially, the result of the interactions between RGR, SLA and LNCa, as well as reflecting intrinsic differences in the efficiency of nitrogen use in the growth process.     

Relative growth rate variation of evergreen and deciduous savanna tree species is driven by different traits

Background and Aims

Plant relative growth rate (RGR) depends on biomass allocation to leaves (leaf mass fraction, LMF), efficient construction of leaf surface area (specific leaf area, SLA) and biomass growth per unit leaf area (net assimilation rate, NAR). Functional groups of species may differ in any of these traits, potentially resulting in (1) differences in mean RGR of groups, and (2) differences in the traits driving RGR variation within each group. We tested these predictions by comparing deciduous and evergreen savanna trees.

Methods

RGR, changes to biomass allocation and leaf morphology, and root non-structural carbohydrate reserves were evaluated for juveniles of 51 savanna species (34 deciduous, 17 evergreen) grown in a common garden experiment. It was anticipated that drivers of RGR would differ between leaf habit groups because deciduous species have to allocate carbohydrates to storage in roots to be able to flush leaves again, which directly compromises their LMF, whereas evergreen species are not subject to this constraint.

Key Results

Evergreen species had greater LMF and RGR than deciduous species. Among deciduous species LMF explained 27 % of RGR variation (SLA 34 % and NAR 29 %), whereas among evergreen species LMF explained between 2 and 17 % of RGR variation (SLA 32–35 % and NAR 38–62 %). RGR and LMF were (negatively) related to carbohydrate storage only among deciduous species.

Conclusions

Trade-offs between investment in carbohydrate reserves and growth occurred only among deciduous species, leading to differences in relative contribution made by the underlying components of RGR between the leaf habit groups. The results suggest that differences in drivers of RGR occur among savanna species because these have different selected strategies for coping with fire disturbance in savannas. It is expected that variation in the drivers of RGR will be found in other functional types that respond differently to particular disturbances.     

Differences in seedling growth behaviour among species: trait correlations across species, and trait shifts along nutrient compared to rainfall gradients

1 Species-pairs from woody dicot lineages were chosen as phylogenetically independent contrasts (PICs) to represent evolutionary divergences along gradients of rainfall and nutrient stress, and within particular habitat types, in New South Wales, Australia. Seedlings were grown under controlled, favourable conditions and measurements were made for various growth, morphological and allocation traits.
2 Trait correlations across all species were identified, particularly with respect to seedling relative growth rate (RGR) and specific leaf area (SLA), a fundamental measure of allocation strategy that reflects the light-capture area deployed per unit of photosynthate invested in leaves.
3 Across all species, SLA, specific root length (SRL) and seed reserve mass were the strongest predictors of seedling RGR. That is, a syndrome of leaf and root surface maximization and low seed mass was typical of high RGR plants. This may be a high-risk strategy for individual seedlings, but one presumably mitigated by a larger number of seedlings being produced, increasing the chance that at least one will find itself in a favourable situation.
4 Syndromes of repeated attribute divergence were identified in the two sets of gradient PICs. Species from lower resource habitats generally had lower SLA. Thus, in this important respect the two gradients appeared to be variants of a more general 'stress' gradient.
5 However, trends in biomass allocation, tissue density, root morphology and seed reserve mass differed between gradients. While SLA and RGR tended to shift together along gradients and in within-habitat PICs, no single attribute emerged as the common, primary factor driving RGR divergences within contrasts. Within-habitat attribute shifts were of similar magnitude to those along gradients.     

Germination and seedling growth of bog plants in relation to the recolonization of milled peatlands

Two controlled experiments were conducted to evaluate the potential for vascular plants to germinate and establish in milled peatlands and to assess whether easily measured plant traits can be used to predict their probable success. Study species included twenty species of perennial herbs, shrubs and trees occurring frequently in undisturbed bogs or abandoned milled bogs in Québec, Canada. First, a glasshouse experiment was performed to test the effect of burial under peat on germination and seedling emergence. Second, a growth chamber experiment was conducted to measure relative growth rate and other growth parameters of seedlings between 1 and 3 weeks of age. In the burial experiment, seedling emergence decreased exponentially with peat depth for most species examined. The slope of the exponential decline varied between species and was strongly correlated to seed mass. Seeds less than 0.1 mg in mass were most sensitive to burial. In the seedling growth experiments, Betula species had the highest absolute and relative growth rates, which may help to explain their prevalence in milled bogs. Relative growth rate (RGR) was not correlated with seed mass, however it was strongly correlated with leaf area ratio (LAR) and especially specific leaf area (SLA) of seedlings, except for species with seed mass less than 0.01 mg. Screening of species for seed mass and SLA should help predict their germination and establishment success or failure in milled peatlands and allow more directed interventions to favour the establishment of desirable species in milled bogs. This revised version was published online in August 2006 with corrections to the Cover Date.     

Phenotypic plasticity and growth temperature: understanding interspecific variability

The subject of this review is the impact of long-term changes in temperature on plant growth and its underlying components. The discussion highlights the extent to which thermal acclimation of metabolism is intrinsically linked to the plasticity of a range of biochemical and morphological traits. The fact that there is often a trade-off between temperature-mediated changes in net assimilation rates (NAR) and biomass allocation [in particular the specific leaf area (SLA)] when plants are grown at different temperatures is also highlighted. Also discussed is the role of temperature-mediated changes in photosynthesis and respiration in determining NAR values. It is shown that in comparisons that do not take phylogeny into account, fast-growing species exhibit greater temperature-dependent changes in RGR, SLA, and NAR than slow-growing plants. For RGR and NAR, such trends are maintained within phylogenetically independent contrasts (i.e. species adapted to more-favourable habitats consistently exhibit greater temperature-mediated changes than their congeneric counterparts adapted to less-favourable habitats). By contrast, SLA was not consistently more thermally plastic in species from favourable habitats. Interestingly, biomass allocation between leaves and roots was consistently more plastic in slow-growing species within individual phylogenetically independent contrasts, when plants were grown under contrasting temperatures. Finally, how interspecific variations in NAR account for an increasing proportion of variability in RGR as growth temperatures decrease is highlighted. Conversely, SLA played a more dominant role in determining interspecific variability in RGR at higher growth temperatures; thus, the importance of SLA in determining interspecific variation in RGR could potentially increase if annual mean temperatures increase in the future.     

Variability in water use efficiency at the leaf level among Mediterranean plants with different growth forms

Assessing natural variability of leaf water use efficiency in plants adapted to extreme conditions of the Mediterranean climate represents an important step in the evaluation of the usefulness of some plant ecophysiological traits under water stress. Eleven Mediterranean species naturally inhabiting the Balearic Islands and corresponding to different growth forms (herbs, semi-deciduous shrubs, woody evergreen shrubs and woody evergreen semi-shrubs) were subject to progressive soil water depletion. Leaf intrinsic water use efficiency was measured by gas exchange at four different degrees of water stress. Under well watered conditions, differences in leaf intrinsic water use efficiency (A _N/g _s) among growth forms were limited to woody evergreen semi-shrubs, which presented the highest values. Under water stress conditions, differences became more evident, with a trend for an increase in A _N/g _s from woody evergreen shrubs, through semi-deciduous shrubs and herbaceous to woody evergreen semi-shrubs. The observed variation in A _N/g _s correlated with several physiological (leaf water potential, soil to leaf hydraulic conductance and stomatal conductance) and morphological (stomatal density) parameters, displaying a general relationship for all growth forms. This suggests that the capacity for withstanding water limitation is adaptive for all Mediterranean species. However, when A _N/g _s was related to leaf mass area, this relationship was not generally applicable, and depended on growth forms, suggesting that different growth forms display specific morphological adjustments in response to water shortage.     

Foliar nutrients in relation to growth, allocation and leaf traits in seedlings of a wide range of woody plant species and types

This study aimed to identify functional correlates of seedling leaf nutrient content among woody species and to characterise functional species groups with respect to leaf nutrient attributes. Seedlings of 81 woody species from the temperate zone of western Europe were grown in a standard laboratory environment with standard, near-optimal nutrient availability. Weight-based leaf N content (N_wght) was positively correlated with mean relative growth rate (RGR), but the correlation with mean RGR was tighter when leaf N was expressed on a whole-plant weight basis: leaf nitrogen weight ratio (LNWR). Area-based leaf N content (N_area) was not associated with mean RGR, but was closely correlated with the quotient of saturated leaf weight and leaf area. Weight-based leaf K content (K_wght) was a close correlate of the saturated/dry weight ratio of the foliage. Within the lower range, K_wght corresponded with growth-related nutrient attributes, but higher values appeared to indicate succulence or remobilisable stored water. Functional groups of species and genera could be distinguished with respect to seedling leaf nutrient attributes. Deciduous woody climbers and scramblers had consistently higher leaf N_wght, LNWR and (apparently) leaf K_wght than other deciduous species or genera, and shrubs had higher values than trees. These differences seemed due partly to variation in specific leaf area. Evergreens had consistently higher leaf N_area than deciduous plants, but there were no significant differences in weight-based leaf nutrient attributes between these two groups, possibly because of `luxury nutrient consumption' by the slow-growing evergreens. Another functional group was that of the nitrogen-fixing species, which had consistently high innate leaf N_wght compared to non-N-fixers. The ecological significance of the leaf nutrient attributes in this study is discussed by comparing the seedling data with those from field-collected material, and by brief reference to the natural habitats of the species. Received: 22 September 1996 / Accepted: 1 March 1997     

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.     

Growth temperature influences the underlying components of relative growth rate: an investigation using inherently fast- and slow-growing plant species

We examined the effect of growth temperature on the underlying components of growth in a range of inherently fast‐ and slow‐growing plant species. Plants were grown hydroponically at constant 18, 23 and 28 °C. Growth analysis was conducted on 16 contrasting plant species, with whole plant gas exchange being performed on six of the 16 species. Inter‐specific variations in specific leaf area (SLA) were important in determining variations in relative growth rate (RGR) amongst the species at 23 and 28 °C but were not related to variations in RGR at 18 °C. When grown at 18 °C, net assimilation rate (NAR) became more important than SLA for explaining variations in RGR. Variations in whole shoot photosynthesis and carbon concentration could not explain the importance of NAR in determining RGR at the lower temperatures. Rather, variations in the degree to which whole plant respiration per unit leaf area acclimated to the different growth temperatures were responsible. Plants grown at 28 °C used a greater proportion of their daily fixed carbon in respiration than did the 18 and 23 °C‐grown plants. It is concluded that the relative importance of the underlying components of growth are influenced by growth temperature, and the degree of acclimation of respiration is of central importance to the greater role played by NAR in determining variations in RGR at declining growth temperatures.     

Effect of soil nitrogen stress on the relative growth rate of annual and perennial grasses in the Intermountain West

A high relative growth rate (RGR) is thought to be an important trait allowing invasive annual grasses to exploit brief increases in nitrogen (N) supply following disturbance in the Intermountain West. Managing soils for low N availability has been suggested as a strategy that may reduce this growth advantage of annual grasses and facilitate establishment of desirable perennials grasses. The objective of this study was to examine the degree to which soil N availability affects RGR and RGR components of invasive annual and desirable perennial grasses. It was hypothesized that (1) invasive annual grasses would demonstrate a proportionately greater reduction in RGR than perennial grasses as soil N stress increased, and (2) the mechanism by which low N availability decreases RGR of annual and perennial grasses would depend on the severity of N stress, with moderate N stress primarily affecting leaf mass ratio (LMR) and severe N stress primarily affecting net assimilation rate (NAR). Three annual and three perennial grasses were exposed to three levels of N availability. RGR and components of RGR were quantified over four harvests. Moderate N stress reduced RGR by decreasing LMR and severe N stress lowered RGR further by decreasing NAR. However, reduction in RGR components was similar between invasive and natives, and as a consequence, annual grasses did not demonstrate a proportionately greater reduction in RGR than perennials under low N conditions. These results suggest managing soil N will do little to reduce the initial growth advantage of annual grasses. Once perennials establish, traits not captured in this short-term study, such as high tissue longevity and efficient nutrient recycling, may allow them to compete effectively with annuals under low N availability. Nevertheless, if soil N management does not facilitate the initial establishment of perennials in annual grass infested communities, then there is little likelihood that such techniques will provide a long-term benefit to restoration projects in these systems.     

Components of variation in seedling potential relative growth rate: phylogenetically independent contrasts

Variation between species in seedling potential relative growth rate (RGR) is among the most important spectra of plant adaptation. Investigations are reported into the components responsible for this variation, using phylogenetically independent contrasts (PICs). The two species for each PIC were selected to diverge in seed mass at least four-fold, seed mass being a known correlate of RGR. Consistent with previous reports, the main influence on RGR differences between species was leaf area per unit leaf mass (SLA), rather than net assimilation rate per leaf area (NAR_a). The PIC design showed that SLA differences both underpinned old RGR divergences between orders and families, and also were repeatedly responsible for more recent RGR divergences between genera and species.     

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.     

Modelling the relative growth rate as a function of plant nitrogen concentration

The relationship between the relative growth rate (RGR) and the nitrogen concentration of the whole plant (PNC) was analyzed by using experimentally determined relations (1) between the PNC and the fraction of dry matter (LWR) and nitrogen in leaves, (2) between the specific leaf area (SLA) and the leaf nitrogen concentration (LNC) and (3) between the net assimilation rate (NAR) and the LNC on an area basis. A strong dependence of RGR on nitrogen concentration resulted from the increase in NAR, LWR and SLA with increasing PNC. A curvilinear relationship between RGR and PNC gave an optimum curve for nitrogen productivity against PNC.