New seed morphological features in Moehringia L. (Caryophyllaceae) and their taxonomic and ecological significance

Abstract

Relative growth rate (RGR) is a fundamental trait for comparative plant ecology but cannot be measured in situ, leading to problems in interpreting vegetation function. However, the components of RGR (net assimilation rate (NAR), leaf area ratio (LAR), leaf weight ratio (LWR), and specific leaf area (SLA)) can be calculated for wild plants from morphological measurements (leaf area, leaf dry mass, whole plant dry mass), which potentially reflect RGR. Seeds of 19 species from Italian prealpine calcareous grasslands were collected and seedlings were cultivated under controlled conditions. RGR, NAR, LAR, LWR and SLA were analysed. The results demonstrated that RGR was positively correlated with SLA and LAR (p < 0.01). Furthermore, LAR was positively correlated with LWR and negatively with NAR (p < 0.05). Monocotyledons showed significantly higher LAR, LWR and NAR than dicotyledons, as the latter allocated a greater proportion of biomass to stems, but RGR and SLA showed no such phylogenetic constraint. Therefore SLA is the most reliable indicator of RGR in ecological and functional surveys of prealpine calcareous grasslands, and has the additional advantage that it can be measured from leaf material alone. Lower mean RGR and SLA for calcareous grassland species suggests that this vegetation is less likely to recover from the effects of disturbance than meadows and dry meadows.     

Relative growth rate in relation to physiological and morphological traits for northern hardwood tree seedlings: species,light environment and ontogenetic considerations

The influence of ontogeny, light environment and species on relationships of relative growth rate (RGR) to physiological and morphological traits were examined for first-year northern hardwood tree seedlings. Three Betulaceae species (Betula papyrifera, Betula alleghaniensis and Ostrya virginiana) were grown in high and low light and Quercus rubra and Acer saccharum were grown only in high light. Plant traits were determined at four ages: 41, 62, 83 and 104 days after germination. In high light (610 mol m^–2 s^–1 PPFD), across species and ages, RGR was positively related to the proportion of the plant in leaves (leaf weight ratio, LWR; leaf area ratio, LAR), in situ rates of average canopy net photosynthesis (A) per unit mass (A_mass) and per unit area (A_area), and rates of leaf, stem and root respiration. In low light (127 mol m^–2 s^–1 PPFD), RGR was not correlated with A_mass and A_area whereas RGR was positively correlated with LAR, LWR, and rates of root and stem respiration. RGR was negatively correlated with leaf mass per area in both high and low light. Across light levels, relationships of CO₂ exchange and morphological characteristics with RGR were generally weaker than within light environments. Moreover, relationships were weaker for plant parameters containing a leaf area component (leaf mass per area, LAR and A_area), than those that were solely mass-based (respiration rates, LWR and A_mass). Across light environments, parameters incorporating the proportion of the plant in leaves and rates of photosynthesis explained a greater amount of variation in RGR (e.g. LWR^*A_mass, R²=0.64) than did any single parameter related to whole-plant carbon gain. RGR generally declined with age and mass, which were used as scalars of ontogeny. LWR (and LAR) also declined for seven of the eight species-light treatments and A declined in four of the five species in high light. Decreasing LWR and A with ontogeny may have been partially responsible for decreasing RGR. Declines in RGR were not due to increased respiration resulting from an increase in the proportion of solely respiring tissue (roots and stems). In general, although LWR declined with ontogeny, specific rates of leaf, stem, and root respiration also decreased. The net result was that whole-plant respiration rates per unit leaf mass decreased for all eight treatments. Identifying the major determinants of variation in growth (e.g. LWR^*A_mass) across light environments, species and ontogeny contributes to the establishment of a framework for exploring limits to productivity and the nature of ecological success as measured by growth. The generality of these relationships both across the sources of variation we explored here and across other sources of variation in RGR needs further study.     

Growth rate and biomass partitioning of wildtype and low-gibberellin tomato (Solanum lycopersicum) plants growing at a high and low nitrogen supply

The growth-promoting effects of gibberellins (GAs) on plants are well documented, but a complete growth analysis at the whole plant level on plants with an altered GA biosynthesis has never been reported. In the present work, the relative growth rate (RGR), biomass partitioning and morphological parameters of wildtype (Wt) tomato ( Solanum lycopersicum L. cv. Moneymaker) plants were compared with those of isogenic ( gib ) mutants with a reduced biosynthesis of gibberellins. GA deficiency reduced RGR and specific leaf area (SLA, leaf area per unit leaf mass) and increased the net assimilation rate (NAR, the rate of biomass increment per unit leaf area). Despite the free access to nitrogen in the rooting medium, the low-GA mutants had a much higher root mass ratio (RMR, the root mass per unit plant biomass) than the Wt, suggesting that the mutants were disturbed in their growth response to nitrate supply. The experiment was repeated at a low exponential nitrate supply, which forced all plants to grow at the same low RGR. The persistence of the differences in RMR at low N-supply indicated that the high RMR of the mutants was a direct effect of low GA, which was independent of nitrate supply. Because the low N-supply increased the RMRs of all genotypes to the same extent, the response of RMR to N-supply does not seem to depend on GA. Although many of the traits of the slow growing GA mutants were very similar to those of inherently slow growing plant species from unproductive habitats, gibberellins are unlikely to be a main determinant of a plant's potential RGR.     

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.     

Differences in developmental plasticity and growth rate among drought-resistant and susceptible cultivars of durum wheat (Triticum turgidum L. var. durum)

Understanding how growth and development of durum wheat cultivars respond to drought could provide a basis to develop crop improvement programmes in drought-affected tropical and subtropical countries. A greenhouse experiment was conducted to study the responses of five durum wheat cultivars to moisture stress at different developmental phases. Phenology, total dry matter (TDM), relative growth rate (RGR), leaf area ratio (LAR), net assimilation rate (NAR), leaf weight ratio (LWR), specific leaf area (SLA) and shoot:root ratio were compared. Pre-anthesis moisture stress delayed phenological development, whereas post-anthesis moisture stress accelerated it. TDM accumulation rate was different between drought-resistant and susceptible cultivars. RGR and its components changed with age and moisture availability. Drought-resistant cultivars had a high RGR in favourable periods of the growing season and a low RGR during moisture stress. In contrast, the drought-susceptible cultivar (Po) showed an opposite trend. LAR explained the differences in RGR (r=0.788) best, whereas the relationship between NAR and RGR was not significant. Even though both LWR and SLA were important factors determining the potential growth rate, LWR was of major importance to describe cultivar differences in LAR, and consequently in RGR. The drought-resistant cultivars Omrabi-5 and Boohai showed vigorous root development and/or a low shoot:root ratio. It is concluded that biomass allocation is the major factor explaining variation in RGR among the investigated durum wheat cultivars.     

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.     

The Interrelated Effects of Stage of Development and Seasonal Changes in Light and Temperature on the Components of Growth in Zea mays

Seeds of Orla 266, a double cross hybrid of Zea mays, were sownin pots in the open 3, 6, and 9 weeks before the individualplants were sampled at consecutive weekly intervals from theend of August. On each occasion the weights of the whole plantand its parts plus leaf area were recorded, and the proceduresof growth analysis applied to the data. Between the youngest and oldest plants over all occasions thegreatest reductions with age were for the rate of leaf expansionand the ratio of leaf area to leaf weight. The net assimilationrate (NAR) and the leaf weight ratio (LWR) were depressed leastwhile the relative growth-rate (RGR) and leaf area ratio (LAR)occupied an intermediate position. The age effects were mostpronounced for RGR, NAR and the ratio of leaf area to leaf weightat the beginning of the experiment, whereas for LAR and LWRthe divergencies were largest at the end. On the basis of the recorded changes in solar radiation andtemperature and prior studies of the interacting effects oflight and temperature on the vegetative growth of Zea it waspossible to predict for the several growth components the patternsof change which should be expected. For the youngest and intermediatepopulations there was close agreement between the expected andobserved values for RGR, NAR, and LAR and it was concluded thatthe time courses were primarily dependent on environmental factors.On the other hand, for the oldest plants from the third intervalonwards the observed values for NAR and less so for RGR werematerially above expectation. It was noted that this divergencecoincided with the rapid extension of the shoot which was confinedto the oldest plants. It is advanced that the higher NAR isassociated with the vertical separation of the leaves.     

Net assimilation rate and relative nitrogen assimilation rate in relation to the dry matter production of alfalfa cultivars

Summary Twelve alfalfa cultivars inoculated with an indigenous strain (RM9) ofRhizobium meliloti, were compared for their seedling morphological characters, and growth characters, including net assimilation rate (NAR), relative growth rate (RGR), leaf area ratio (LAR) and relative nitrogen assimilation rate (R_N). Highly significant differences were obtained between cultivars for most characters.Simple correlation showed that NAR influenced RGR (r=0.91) more than leaf area ratio (LAR) (r=–0.44), and that most characters measured were highly correlated with seedling dry weight. Factor analysis showed that NAR, RGR and R_N contributed 25% of the total variation in the dependence structure. The grouping indicated that the higher the NAR and R_N the greater was the RGR. Path-coefficient analysis showed that NAR had more important direct and indirect effects than R_N in dry matter accumulation. The relationship implied that selection for plants with high NAR, or high efficiency in converting light energy to dry matter production could contribute greater N₂ fixation in alfalfa.     

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.     

Close relationship between leaf life span and seedling relative growth rate in temperate hardwood species

The life span of resource-acquiring organs (leaves, shoots, fine roots) is closely associated with species successional position and environmental resource availability. We examined to what extent leaf life span is related to inter- and intraspecific variation in seedling relative growth rate (RGR). We examined relationships between relative growth rate in mass (RGR_M) or height (RGR_H) and leaf life span, together with classical RGR_M components [net assimilation rate (NAR), specific leaf area (SLA), leaf weight ratio (LWR), and leaf area ratio (LAR)] for seedlings of five hardwood species of different successional position across a wide range of environmental resource availability, including the presence or absence of leaf litter in shaded forest understory, small canopy gaps, and large canopy gaps. Both SLA and LAR were negatively correlated with RGR_M along the environmental gradient for all species. However, positive correlations were observed among species within microsites, indicating that these two components cannot consistently explain the variation in RGR_M. Both NAR and LWR affect interspecific, but not intraspecific, variation in RGR_M. Leaf life span was negatively correlated with either RGR_M or RGR_H in both inter- and intraspecific comparisons. Species with short-lived, physiologically active leaves have high growth rates, particularly in resource-rich environments. Consequently, leaf life span is a good predictor of seedling RGR. Leaf life span affects plant performance and has a strong and consistent effect on tree seedling growth, even among contrasting environments.     

Seedling growth of five tropical dry forest tree species in relation to light and nitrogen gradients

Aims Increasing anthropogenic nitrogen (N) deposition has been claimed to induce changes in species composition and community dynamics. A greenhouse experiment was conducted to examine the effect of increased N availability on growth and functional attributes of seedlings of five tree species with different life history characteristics under varying irradiances. The following questions have been addressed: (i) how do the pioneer and non-pioneer species respond in absolute growth and relative growth rate (RGR) to the interaction of light and nitrogen? (ii) how does the interaction between irradiance and nitrogen availability modulate growth attributes (i.e. functional attributes)? (iii) is there any variation in growth responses between leguminous and non-leguminous species along the light and nitrogen gradients?Methods Seedlings of five tree species (Acacia catechu, Bridelia retusa, Dalbergia sissoo, Lagerstroemia parviflora and Terminalia arjuna) were subjected to twelve combinations of irradiance and N levels. Various growth traits, including height (HT), basal area (BA), whole plant dry biomass (M D), leaf mass per unit area (LMA), leaf area ratio (LAR), net assimilation rate (NAR), RGR, biomass fractions, root-to-shoot ratio (R:S) and leaf nitrogen content, were studied to analyse intra- and inter-specific responses to interacting light and N gradients.Important findings Significant interactions for irradiance and N availability for majority of growth attributes indicates that growth and biomass allocation of seedlings were more responsive to N availability under high irradiance. However, species responded differentially to N addition and they did not follow successional status. Slow growers (B. retusa, a shade-tolerant species and L. parviflora, a light demander) exhibited greater response to N enrichment than the fast growers (A. catechu, D. sissoo and T. arjuna). However, N-mediated increment in growth traits was greater in non-legumes (B. retusa, L. parviflora and T. arjuna) compared with that of legumes (A. catechu and D. sissoo). Allocation of biomass to root was strongly suppressed at the highest N supply across species; however, at high irradiance and high N availability, a greater suppression in R:S ratio was observed for B. retusa. NAR was a stronger determinant of RGR relative to LAR, suggesting its prominent role in increased RGR along increasing irradiances. Overall, a higher growth response of slow-growing species to elevated N levels, particularly the non-pioneers (B. retusa and L. parviflora) suggests that future N deposition may lead to perturbations in competition hierarchies and species composition, ultimately affecting community dynamics in nutrient-poor tropical dry forests.     

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.     

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.     

Seedling growth strategies in Bauhinia species: comparing lianas and trees

BACKGROUND AND AIMS: Lianas are expected to differ from trees in their growth strategies. As a result these two groups of woody species will have different spatial distributions: lianas are more common in high light environments. This study determines the differences in growth patterns, biomass allocation and leaf traits in five closely related liana and tree species of the genus Bauhinia. METHODS: Seedlings of two light-demanding lianas (Bauhinia tenuiflora and B. claviflora), one shade-tolerant liana (B. aurea), and two light-demanding trees (B. purpurea and B. monandra) were grown in a shadehouse at 25% of full sunlight. A range of physiological, morphological and biomass parameters at the leaf and whole plant level were compared among these five species. KEY RESULTS: The two light-demanding liana species had higher relative growth rate (RGR), allocated more biomass to leaf production [higher leaf mass fraction (LMF) and higher leaf area ratio (LAR)] and stem mass fraction (SMF), and less biomass to the roots [root mass fraction (RMF)] than the two tree species. The shade-tolerant liana had the lowest RGR of all five species, and had a higher RMF, lower SMF and similar LMF than the two light-demanding liana species. The two light-demanding lianas had lower photosynthetic rates per unit area (A(area)) and similar photosynthetic rates per unit mass (A(mass)) than the trees. Across species, RGR was positively related to SLA, but not to LAR and A(area). CONCLUSIONS: It is concluded that the faster growth of light-demanding lianas compared with light-demanding trees is based on morphological parameters (SLA, LMF and LAR), and cannot be attributed to higher photosynthetic rates at the leaf level. The shade-tolerant liana exhibited a slow-growth strategy, compared with the light-demanding species.     

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.     

Applicability and limitations of optimal biomass allocation models: a test of two species from fertile and infertile habitats

BACKGROUND AND AIMS: The practical applicability of optimal biomass allocation models is not clear. Plants may have constraints in the plasticity of their root : leaf ratio that prevent them from regulating their root : leaf ratio in the optimal manner predicted by the models. The aim of this study was to examine the applicability and limitations of optimal biomass allocation models and to test the assumption that regulation of the root : leaf ratio enables maximization of the relative growth rate (RGR). METHODS: Polygonum cuspidatum from an infertile habitat and Chenopodium album from a fertile habitat were grown under a range of nitrogen availabilities. The biomass allocation, leaf nitrogen concentration (LNC), RGR, net assimilation rate (NAR), and leaf area ratio (LAR) of each species were compared with optimal values determined using an optimal biomass allocation model. KEY RESULTS: The root : leaf ratio of C. album was smaller than the optimal ratio in the low-nitrogen treatment, while it was almost optimal in the high-nitrogen treatment. In contrast, the root : leaf ratio of P. cuspidatum was close to the optimum under both high- and low-nitrogen conditions. Owing to the optimal regulation of the root : leaf ratio, C. album in the high-nitrogen treatment and P. cuspidatum in both treatments had LNC and RGR (with its two components, NAR and LAR) close to their optima. However, in the low-nitrogen treatment, the suboptimal root : leaf ratio of C. album led to a smaller LNC than the optimum, which in turn resulted in a smaller NAR than the optimum and RGR than the theoretical maximum RGR. CONCLUSIONS: The applicability of optimal biomass allocation models is fairly high, although constraints in the plasticity of biomass allocation could prevent optimal regulation of the root : leaf ratio in some species. The assumption that regulation of the root : leaf ratio enables maximization of RGR was supported.     

Relative contributions of leaf area ratio and net assimilation rate to change in growth rate depend on growth temperature: comparative analysis of subantarctic and alpine grasses

The present study shows that the relative contributions of leaf area ratio (LAR) and net assimilation rate (NAR) to variation among species in relative growth rate (RGR) depend on growth temperature. We grew three subantarctic and three alpine Poa species at daytime temperatures of 7, 12 and 17 degrees C, and analysed interspecific and temperature-related variation in RGRs by growth analysis. Variation in NAR accounted for most of the interspecific differences in RGR at low growth temperature, whereas variation in both NAR and LAR contributed strongly to interspecific differences in RGR at high growth temperature. For most species, the increase in RGR from 7 to 12 degrees C was attributable to an increase in LAR, whereas the increase in RGR from 12 to 17 degrees C was attributable to an increase in NAR. There were no differences between native subantarctic and alpine species in the plasticity of growth responses to temperature. However, Poa annua, a species introduced to the subantarctic, showed much greater growth plasticity than other species. There was little difference among species in tolerance of high-temperature extremes.     

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.     

Roles of succession,light, nutrients and disturbance on population vigor and maintenance of the rare plant Solidago shortii (Asteraceae)

Results of field and glasshouse experiments on Solidago shortii, and our observations on this species over many years, were used to construct a conceptual model of the roles of succession, light, soil nutrients and disturbance on population vigor and maintenance of this federal-endangered species. As cover of woody vegetation increased at a population site between 1986 and 1992, number of flowering ramets of S. shortii significantly decreased but number of vegetative ramets remained nearly constant. Adult plants transplanted into a redcedar thicket and those shaded in a glasshouse produced many fewer flowering ramets and capitula per flowering ramet and less biomass and had higher mortality than those in the open. Seedlings/juveniles shaded in a glasshouse had significantly less dry biomass; lower RGR, NAR, leaf area and root/shoot ratio and higher LAR, SLA and LWR than nonshaded ones. In a field site and glasshouse, fertilized plants (NPK) consistently had more flowering ramets and capitula per flowering ramet than nonfertilized ones. Hierarchy of dry weight of plants grown in a glasshouse in soils derived from five types of bedrock was phosphatic limestone > calcareous shale > sandstone > black shale = dolomite. Flowering and biomass production in the field-fertilizer and soil-type experiments were associated closely with levels of P. Number of flowering ramets significantly increased in plants transferred from shaded to nonshaded glasshouse conditions, but no such increase occurred after opening the canopy above plants in a thicket. Both high light and high nutrient levels apparently are necessary to maintain high vigor of S. shortii. In areas subject to invasion by woody plants, periodic high intensity disturbance may be required to prevent population extirpation.     

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).