Plasticity In Specific Leaf Area Research Articles

Selection favors adaptive plasticity in a long-term reciprocal transplant experiment.

Spatial and temporal environmental variation can favor the evolution of adaptive phenotypic plasticity, such that genotypes alter their phenotypes in response to local conditions to maintain fitness across heterogeneous landscapes. When individuals show greater fitness in one habitat than another, asymmetric migration can restrict adaptation to the lower quality environment. In these cases, selection is predicted to favor traits that enhance fitness in the higher-quality (source) habitat at the expense of fitness in the marginal (sink) habitat.Here, we test whether plasticity is adaptive in a system regulated by demographic source-sink dynamics.Vaccinium elliottii (Ericaceae) occurs in dry upland and flood-prone bottomland forests throughout the southeastern United States, but has larger populations and higher average individual fitness in upland sites. We conducted a multi-year field experiment to evaluate whether plasticity in foliar morphology increases survival and lifespan. Both across and within habitats, selection favored plasticity in specific leaf area, stomatal density, and leaf size. Stabilizing selection acted on plasticity in stomatal density within habitats, suggesting that extreme levels of plasticity are disadvantageous. Thus, even in systems driven by source-sink dynamics, temporal and spatial variation in conditions across the landscape and within habitat types can favor the evolution of plasticity.

Traits and their plasticity determine responses of plant performance and community functional property to nitrogen enrichment in a boreal peatland

Nitrogen (N) enrichment is well-known to cause declines in plant diversity, yet it remains a challenge to predict which species would be winners or losers. Here, we examined whether traits and their plasticity could predict plant performance under N enrichment, and how changes in plant performance and trait expression could affect community functional property. Relative cover, six leaf traits, and maximum plant height of eight common vascular plants were measured after six-year N addition (0, 3, 6, 12 g N m−2 year−1) in a boreal peatland in Northeast China. Plant performance responding to N enrichment and trait plasticity were separately calculated as changes in relative cover and trait values between control and N addition treatments. Moreover, we quantified relative contributions of species turnover and intraspecific trait variation to changes in functional property. Changes in plant performance were significantly related to plasticity in specific leaf area (SLA) and leaf N concentration (LN) under low N addition, but to maximum plant height under high N addition. When scaled up to the community level, intraspecific trait variation significantly contributed to changes in community-weighted mean leaf area, SLA, leaf tissue density, and LN, and functional diversity of all six leaf traits. These findings highlight the significance of trait plasticity in predicting responses of plant performance and community functional property to N enrichment, and suggest that trait-based approaches are useful in predicting ecological consequences of environmental changes in boreal peatlands.

Plasticity of functional traits of tree of heaven is higher in exotic than in native habitats

Novel features related to turgor loss adjustment and higher ability to modulate leaf-construction costs could improve A. altissima fitness in its invaded range. Phenotypic plasticity has been recently proposed to play an important role in invasion process. More precisely, phenotypic plasticity of alien invasive species could be higher in their exotic rather than native habitat, probably because of the release from biotic or abiotic constraints. In this paper, several plant functional traits were analysed on Ailanthus altissima (Mill.) Swingle, a highly invasive species in Europe, aiming at providing a comparison among key functional traits measured on individuals growing in their native habitat and to test if values of functional traits and their related plasticity are higher in exotic habitats. Our analysis pinpointed that variability of functional traits in the native habitat was mainly driven by different irradiance, temperature and evaporative demand of the studied sites, in accordance to the traits trade-offs in the Leaf Economic Spectrum. Physiological traits related to drought tolerance were different between native and exotic habitats. In the native one, A. altissima relied on osmoregulation processes to adjust leaf turgor loss point (Ψtlp), as the osmotic potential at full turgor (π0) were lower in drier sites. In the exotic habitat, individuals in drier sites had similar π0 but lower wall elasticity (e), suggesting that leaves had thinner cell walls. Moreover, plasticity in specific leaf area and e were higher in the exotic habitat. The novel features related to Ψtlp adjustment and the higher plasticity in traits related to leaf-construction costs could increase A. altissima fitness in response to different environmental conditions in its invasive range.

Does greater specific leaf area plasticity help plants to maintain a high performance when shaded?

It is frequently assumed that phenotypic plasticity can be very advantageous for plants, because it may increase environmental tolerance (fitness homeostasis). This should, however, only hold for plastic responses that are adaptive, i.e. increase fitness. Numerous studies have shown shade-induced increases in specific leaf area (SLA), and there is wide consensus that this plastic response optimizes light capture and thus has to be adaptive. However, it has rarely been tested whether this is really the case. In order to identify whether SLA plasticity does contribute to the maintenance of high biomass of plant species under shaded conditions, a meta-analytical approach was employed. The data set included 280 species and 467 individual studies from 32 publications and two unpublished experiments. Plants increased their SLA by 55·4% on average when shaded, while they decreased their biomass by 59·9%. Species with a high SLA under high-light control conditions showed a significantly greater ability to maintain biomass production under shade overall. However, in contrast to the expectation of a positive relationship between SLA plasticity and maintenance of plant biomass, the results indicated that species with greater SLA plasticity were less able to maintain biomass under shade. Although a high SLA per se contributes to biomass homeostasis, there was no evidence that plasticity in SLA contributes to this. Therefore, it is argued that some of the plastic changes that are frequently thought to be adaptive might simply reflect passive responses to the environment, or result as by-products of adaptive plastic responses in other traits.

Invasive Eupatorium catarium and Ageratum conyzoides benefit more than does a common native plant from nutrient addition in both competitive and non‐competitive environments

AbstractWe explored the roles of phenotypic plasticity and competition in contributing to the success of Eupatorium catarium, a recently invasive noxious annual forb in South China. We compared the growth and functional traits of E. catarium with those of a common native plant, Vernonia cinerea, and a long‐term historic invader, Ageratum conyzoides, along a nutrient gradient under both competitive and non‐competitive conditions. Under non‐competitive conditions, biomass differed little between species under low‐nutrient conditions. However, nutrient addition resulted in a greater increase in biomass of the two invasive species than that of V. cinerea. The greater increase in biomass in the two invasive plants may be partly explained by their greater plasticity in specific leaf area. The competition experiments involved different combinations of species: plants of E. catarium were grown with either a plant of A. conyzoides or with a plant of V. cinerea, or individual plants of all three species were grown in competition with a grassland sward grown from 1.5 g seeds of Digitaria radicosa. Relative to non‐competitive conditions, competition further increased the biomass difference between the two invasive plants and V. cinerea under high nutrient conditions. A. conyzoides outcompeted E. catarium in terms of biomass production under high nutrient conditions. Overall, these results indicate that A. conyzoides and E. catarium tend to outperform V. cinerea under high nutrient conditions and in both competitive and non‐competitive environments. However, E. catarium cannot outperform A. conyzoides in terms of biomass production under each of the nutrient and competitive conditions.

Contrasting effects of intraspecific trait variation on trait-based niches and performance of legumes in plant mixtures.

Niche differentiation, assumed to be a key mechanism of species coexistence, requires that species differ in their functional traits. So far it remains unclear to which extent trait plasticity leads to niche shifts of species at higher plant diversity, thereby increasing or decreasing niche overlap between species. To analyse this question it is convenient to measure niches indirectly via the variation in resource-uptake traits rather than directly via the resources used. We provisionally call these indirectly measured niches trait-based niches. We studied shoot- and leaf-morphological characteristics in seven legume species in monoculture and multi-species mixture in experimental grassland. Legume species varied in the extent of trait variation in response to plant diversity. Trait plasticity led to significant shifts in species niches in multiple dimensions. Single-species niches in several traits associated with height growth and filling of canopy space were expanded, while other niche dimensions were compressed or did not change with plant diversity. Niche separation among legumes decreased in dimensions related to height growth and space filling, but increased in dimensions related to leaf size and morphology. The total extent of occupied niche space was larger in mixture than in the combined monocultures for dimensions related to leaf morphology and smaller for dimensions related to whole-plant architecture. Taller growth, greater space filling and greater plasticity in shoot height were positively, while larger values and greater plasticity in specific leaf area were negatively related with increased performance of species in mixture. Our study shows that trait variation in response to plant diversity shifts species niches along trait axes. Plastically increased niche differentiation is restricted to niche dimensions that are apparently not related to size-dependent differences between species, but functional equivalence (convergence in height growth) rather than complementarity (divergence in traits associated with light acquisition) explains increased performance of legumes in mixture.

Within-canopy variation in photosynthetic capacity, SLA and foliar N in temperate broad-leaved trees with contrasting shade tolerance

The relative shade tolerance of T. cordata , F. sylvatica , and C. betulus in mature stands is based on different species-specific carbon and nitrogen allocation patterns. The leaf morphology and photosynthetic capacity of trees are remarkably plastic in response to intra-canopy light gradients. While most studies examined seedlings, it is not well understood how plasticity differs in mature trees among species with contrasting shade tolerance. We studied light-saturated net photosynthesis (A max), maximum carboxylation rate (V cmax), electron transport capacity (J max) and leaf dark respiration (R d) along natural light gradients in the canopies of 26 adult trees of five broad-leaved tree species in a mixed temperate old-growth forest (Fraxinus excelsior, Acer pseudoplatanus, Carpinus betulus, Tilia cordata and Fagus sylvatica), representing a sequence from moderately light-demanding to highly shade-tolerant species. We searched for species differences in the dependence of photosynthetic capacity on relative irradiance (RI), specific leaf area (SLA) and nitrogen per leaf area (N a ). The three shade-tolerant species (C. betulus, T. cordata, F. sylvatica) differed from the two more light-demanding species by the formation of shade leaves with particularly high SLA but relatively low N a and consequently lower area-based A max, and a generally higher leaf morphological and functional plasticity across the canopy. Sun leaf morphology and physiology were more similar among the two groups. The three shade-tolerant species differed in their shade acclimation strategies which are primarily determined by the species’ plasticity in SLA. Under low light, T. cordata and F. sylvatica increased SLA, mass-based foliar N and leaf size, while C. betulus increased solely SLA exhibiting only low intra-crown plasticity in leaf morphology and N allocation patterns. This study with mature trees adds to our understanding of tree species differences in shade acclimation strategies under the natural conditions of a mixed old-growth forest.

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

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. 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. 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.

PLASTICITY OF PHYSIOOLGY INLOBELIA: TESTING FOR ADAPTATION AND CONSTRAINT

Phenotypic plasticity is thought to be a major mechanism allowing sessile organisms such as plants to adapt to environmental heterogeneity. However, the adaptive value of many common plastic responses has not been tested by linking these responses to fitness. Even when plasticity is adaptive, costs of plasticity, such as the energy necessary to maintain regulatory pathways for plastic responses, may constrain its evolution. We used a greenhouse experiment to test whether plastic physiological responses to soil water availability (wet vs. dry conditions) were adaptive and/or costly in the congeneric wildflowers Lobelia cardinalis and L. siphilitica. Eight physiological traits related to carbon and water uptake were measured. Specific leaf area (SLA), photosynthetic rate (A), stomatal conductance (gs), and photosynthetic capacity (Amax) responded plastically to soil water availability in L. cardinalis. Plasticity in Amax was maladaptive, plasticity in A and g(s) was adaptive, and plasticity in SLA was adaptively neutral. The nature of adaptive plasticity in L. cardinalis, however, differed from previous studies. Lobelia cardinalis plants with more conservative water use, characterized by lower g(s), did not have higher fitness under drought conditions. Instead, well-watered L. cardinalis that had higher g(s) had higher fitness. Only Amax responded plastically to drought in L. siphilitica, and this response was adaptively neutral. We detected no costs of plasticity for any physiological trait in either L. cardinalis or L. siphilitica, suggesting that the evolution of plasticity in these traits would not be constrained by costs. Physiological responses to drought in plants are presumed to be adaptive, but our data suggest that much of this plasticity can be adaptively neutral or maladaptive.

PLASTICITY OF PHYSIOLOGY IN LOBELIA: TESTING FOR ADAPTATION AND CONSTRAINT

Phenotypic plasticity is thought to be a major mechanism allowing sessile organisms such as plants to adapt to environmental heterogeneity. However, the adaptive value of many common plastic responses has not been tested by linking these responses to fitness. Even when plasticity is adaptive, costs of plasticity, such as the energy necessary to maintain regulatory pathways for plastic responses, may constrain its evolution. We used a greenhouse experiment to test whether plastic physiological responses to soil water availability (wet vs. dry conditions) were adaptive and/or costly in the congeneric wildflowers Lobelia cardinalis and L. siphilitica. Eight physiological traits related to carbon and water uptake were measured. Specific leaf area (SLA), photosynthetic rate (A), stomatal conductance (gs), and photosynthetic capacity (Amax) responded plastically to soil water availability in L. cardinalis. Plasticity in Amax was maladaptive, plasticity in A and gs was adaptive, and plasticity in SLA was adaptively neutral. The nature of adaptive plasticity in L. cardinalis, however, differed from previous studies. Lobelia cardinalis plants with more conservative water use, characterized by lower gs, did not have higher fitness under drought conditions. Instead, well-watered L. cardinalis that had higher gs had higher fitness. Only Amax responded plastically to drought in L. siphilitica, and this response was adaptively neutral. We detected no costs of plasticity for any physiological trait in either L. cardinalis or L. siphilitica, suggesting that the evolution of plasticity in these traits would not be constrained by costs. Physiological responses to drought in plants are presumed to be adaptive, but our data suggest that much of this plasticity can be adaptively neutral or maladaptive.

Ecophysiological responses to light availability in three Blechnum species (Pteridophyta, Blechnaceae) of different ecological breadth

In Chilean evergreen temperate forest, fern species of the genus Blechnum occur in diverse microhabitats ranging from large gaps to heavily shaded understoreys. We hypothesised that differences in the ecological breadth of three co-occurring Blechnum species would be associated with differences in magnitude of ecophysiological responses to light availability. We quantified the field distribution of each species in relation to diffuse light availability (% canopy openness), and measured in situ variation in photosynthetic capacity (A), dark respiration (R (d)) and specific leaf area (SLA) across the light gradient. The response of SLA of each species was also evaluated in a common garden in two light conditions (understorey and forest edge). The three Blechnum species differed significantly in the range of light environments occupied (breadth: B. chilense > B. hastatum > B. mochaenum). Despite significant interspecific differences in average A and R (d), the response of these traits to light availability did not differ among species. However, there was significant interspecific variation in both the mean value and the plasticity of SLA to light availability, the species with least ecological breadth (B. mochaenum) showing a flatter reaction norm (lower response) than its two congeners. This pattern was also found in the common garden experiment. The adjustment of leaf morphology (SLA) to light availability appears to be an important mechanism of acclimation in these Blechnum species. The narrow range of light environments occupied by B. mochaenum may be at least partly attributable to its inability to display phenotypic plasticity in SLA to changes in light availability.

Compensatory responses of late watergrass (Echinochloa phyllopogon) and rice to resource limitations

The development of optimal weed management strategies that rely, in part, on crop interference will require an understanding of how weeds compensate for limitations in above- and belowground resources. Trade-offs in the leaf morphology and biomass partitioning of rice and late watergrass were investigated under glasshouse conditions in 1999 and 2000. Both species responded to shade with increased height, reduced biomass, greater partitioning of biomass to leaves, and greater leaf area ratios. At the lowest light level (18% sunlight), plants of both species showed little response to nitrogen (N). However, height, tillers, biomass, and leaf area increased for plants grown at 50% and full sunlight as N increased from 0 to 224 kg N ha−1. Late watergrass exhibited more plasticity in specific leaf area and root weight ratio than rice in response to shade. This plasticity contributed to the ability of late watergrass to maintain a higher percent of its tillers and total dry weight than rice when sunlight was reduced by 50%. These results support the hypothesis that except at low light levels, limited N further reduces the growth of shaded late watergrass plants. Thus, weed management strategies that limit the plasticity of late watergrass by manipulating light and N availability are likely to be more effective than strategies that rely on manipulating a single resource.

Comparative growth analysis of tropical forest and savanna woody plants using phylogenetically independent contrasts

Summary 1 The savanna–forest boundary in the tropics is marked by a discontinuity not only in tree density, but also in species composition, with few tree species regularly occurring in both savanna and forest environments. 2 We performed comparative growth analysis for nine congeneric pairs, each containing one savanna species and one forest species, grown in a factorial design involving two light and two nutrient levels. 3 Contrary to predictions, there was no difference in relative growth rates (RGR) between savanna and forest species. However, there were clear differences in allocation patterns and in phenotypic responses to light intensity. Savanna species allocated more biomass to roots and maintained lower leaf area per unit plant mass (LAR), and lower leaf area per unit leaf mass (SLA). 4 Savanna species also exhibited greater phenotypic plasticity in specific leaf area, leaf area ratio and net assimilation rate in response to light intensity. An increase in LAR in response to shading of savanna species offset a decrease in net assimilation rate per unit leaf area (NAR), such that RGR was largely unaffected. 5 For most plant traits measured, more of the interspecific variation could be attributed to differences among genera than to differences between the two functional types, indicating that these seedling traits are highly conserved within the congeneric species pairs. Many of these traits were correlated to seed mass, which is itself highly conserved within genera.

Interspecific consistency and intraspecific variability of specific leaf area with respect to irradiance and nutrient availability

Specific leaf area (SLA) is a common variable in comparative plant ecology that is often measured on field-grown plants, yet SLA is known to be a plastic trait. The motivation for this study was therefore to determine the degree to which environmentally induced plasticity in SLA might obscure interspecific trends. Specifically, we tested the hypothesis that SLA is more responsive to nutrient stress at low irradiance than at high irradiance. To test this hypothesis we grew plants of six species of herbaceous Asteraceae in factorial combinations of 500, 250, and 125 µmol m-2 s-1 irradiance and a full strength (1/1), a 1/5, and a 1/10 dilution of a modified Hoagland hydroponic solution. The full strength solution had 8 mM of nitrogen. Plants were harvested at 21, 28, 35, and 42 days from germination. SLA varied significantly between species, harvest dates, and experimental treatments. In general, SLA decreased with age, but more severely at low irradiance. SLA increased with decreasing irradiance. SLA also increased with increasing nutrient concentration, but this effect disappeared in the older plants at the higher irradiance. These trends were not seen in every species. Our hypothesis was supported: SLA is more responsive to nutrient stress at low irradiance than at high irradiance, but the exact response is species-specific. Despite these environmentally induced changes in SLA at the intraspecific level, the interspecific differences were maintained across environments, justifying the use of SLA as a useful trait in comparative plant ecology.