Hygrochastic capsule dehiscence supports safe site strategies in New Zealand alpine Veronica (Plantaginaceae)

Background and Aims

Hygrochasy is a capsule-opening mechanism predominantly associated with plants in arid habitats, where it facilitates spatially and temporally restricted dispersal. Recently, hygrochastic capsules were described in detail for the first time in alpine Veronica in New Zealand. The aim of the present study was to investigate whether hygrochastic capsules are an adaptation of alpine Veronica to achieve directed dispersal to safe sites. We expect that by limiting dispersal to rainfall events, distances travelled by seeds are short and confine them to small habitat patches where both seedlings and adults have a greater chance of survival.

Methods

Dispersal distances of five hygrochastic Veronica were measured under laboratory and field conditions and the seed shadow was analysed. Habitat patch size of hygrochastic Veronica and related non-hygrochastic species were estimated and compared.

Key Results

Dispersal distances achieved by dispersal with raindrops did not exceed 1 m but weather conditions could influence the even distribution of seeds around the parent plant. Compared with related Veronica species, hygrochastic Veronica mostly grow in small, restricted habitat patches surrounded by distinctly different habitats. These habitat patches provide safe sites for seeds due to their microtopography and occurrence of adult cushion plants. Non-hygrochastic Veronica can be predominantly found in large habitats without clearly defined borders and can be spread over long distances along rivers.

Conclusions

The results suggest that hygrochasy is a very effective mechanism of restricting seed dispersal to rainfall events and ensuring short-distance dispersal within a small habitat patch. It appears that it is an adaptation for directed dispersal to safe sites that only exist within the parent habitat.     

Nitrogen and water addition reduce leaf longevity of steppe species

Background and aims

Changes in supplies of resources will modify plant functional traits. However, few experimental studies have addressed the effects of nitrogen and water variations, either singly or in combination, on functional traits.

Methods

A 2-year field experiment was conducted to test the effects of nitrogen and water addition on leaf longevity and other functional traits of the two dominant (Agropyron cristatum and Stipa krylovii) and three most common species (Cleistogenes squarrosa, Melilotoides ruthenica and Potentilla tanacetifolia) in a temperate steppe in northern China.

Key Results

Additional nitrogen and water increased leaf nitrogen content and net photosynthetic rate, and changed other measured functional traits. Leaf longevity decreased significantly with both nitrogen addition (–6 days in 2007 and –5·4 days in 2008; both P < 0·001) and watering (–13 days in 2007 and –9·9 days in 2008; both P < 0·001), and significant differences in leaf longevity were also found among species. Nitrogen and water interacted to affect leaf longevity and other functional traits. Soil water content explained approx. 70 % of the shifts in leaf longevity. Biomass at both species and community level increased under water and nitrogen addition because of the increase in leaf biomass production per individual plant.

Conclusions

The results suggest that additional nitrogen and water supplies reduce plant leaf longevity. Soil water availability might play a fundamental role in determining leaf longevity and other leaf functional traits, and its effects can be modified by soil nitrogen availability in semi-arid areas. The different responses of species to resource alterations may cause different global change ramifications under future climate change scenarios.     

Linking ethylene to nitrogen-dependent leaf longevity of grass species in a temperate steppe

Background and Aims

Leaf longevity is an important plant functional trait that often varies with soil nitrogen supply. Ethylene is a classical plant hormone involved in the control of senescence and abscission, but its role in nitrogen-dependent leaf longevity is largely unknown.

Methods

Pot and field experiments were performed to examine the effects of nitrogen addition on leaf longevity and ethylene production in two dominant plant species, Agropyron cristatum and Stipa krylovii, in a temperate steppe in northern China.

Key Results

Nitrogen addition increased leaf ethylene production and nitrogen concentration but shortened leaf longevity; the addition of cobalt chloride, an ethylene biosynthesis inhibitor, reduced leaf nitrogen concentration and increased leaf longevity. Path analysis indicated that nitrogen addition reduced leaf longevity mainly through altering leaf ethylene production.

Conclusions

These findings provide the first experimental evidence in support of the involvement of ethylene in nitrogen-induced decrease in leaf longevity.     

Plant resource-use strategies: the importance of phenotypic plasticity in response to a productivity gradient for two subalpine species

Background and Aims

Functional traits are indicators of plant interactions with their environment and the resource-use strategies of species can be defined through some key functional traits. The importance of genetic variability and phenotypic plasticity in trait variations in response to a common environmental change was investigated in two subalpine species.

Methods

Two species with contrasted resource-use strategies, Dactylis glomerata and Festuca paniculata, were grown along a productivity gradient in a greenhouse experiment. Functional traits of different genotypes were measured to estimate the relative roles of phenotypic plasticity and genetic variability, and to compare their levels of phenotypic plasticity.

Key Results

Trait variability in the field for the two species is more likely to be the result of phenotypic plasticity rather than of genetic differentiation between populations. The exploitative species D. glomerata expressed an overall higher level of phenotypic plasticity compared with the conservative species F. paniculata. In addition to different amplitudes of phenotypic plasticity, the two species differed in their pattern of response for three functional traits relevant to resource use (specific leaf area, leaf dry matter content and leaf nitrogen content).

Conclusions

Functional trait variability was mainly the result of phenotypic plasticity, with the exploitative species showing greater variability. In addition to average trait values, two species with different resource-use strategies differed in their plastic responses to productivity.     

Interspecific gene flow in a multispecies oak hybrid zone in the Sierra Tarahumara of Mexico

Background and Aims

Interspecific gene flow can occur in many combinations among species within the genus Quercus, but simultaneous hybridization among more than two species has been rarely analysed. The present study addresses the genetic structure and morphological variation in a triple hybrid zone formed by Q. hypoleucoides, Q. scytophylla and Q. sideroxyla in north-western Mexico.

Methods

A total of 247 trees from ten reference and 13 presumed intermediate populations were characterized using leaf shape variation and geometric morphometrics, and seven nuclear microsatellites as genetic markers. Discriminant function analysis was performed for leaf shape variation, and estimates of genetic diversity and structure, and individual Bayesian genetic assignments were obtained.

Key Results

Reference populations formed three completely distinct groups according to discriminant function analysis based on the morphological data, and showed low, but significant, genetic differentiation. Populations from the zone of contact contained individuals morphologically intermediate between pairs of species in different combinations, or even among the three species. The Bayesian admixture analysis found that three main genetic clusters best fitted the data, with good correspondence of reference populations of each species to one of the genetic clusters, but various degrees of admixture evidenced in populations from the contact area.

Conclusions

The three oak species have formed a complex hybrid zone that is geographically structured as a mosaic, and comprising a wide range of genotypes, including hybrids between different species pairs, backcrosses and probable triple hybrids.     

Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub

Background and Aims

Understorey evergreen species commonly have a higher leaf nitrogen content in winter than in summer. Tested here is a hypothesis that such changes in leaf nitrogen content maximize nitrogen-use efficiency, defined as the daily carbon gain per unit nitrogen, under given temperature and irradiance levels.

Methods

The evergreen shrub Aucuba japonica growing naturally at three sites with different irradiance regimes in Japan was studied. Leaf photosynthetic characteristics, Rubisco and leaf nitrogen with measurements of temperature and irradiance monthly at each site were determined. Daily carbon gain was determined as a function of leaf nitrogen content to calculate the optimal leaf nitrogen content that maximized daily nitrogen-use efficiency.

Key Results

As is known, the optimal leaf nitrogen content increased with increasing irradiance. The optimal leaf nitrogen content also increased with decreasing temperature because the photosynthetic capacity per Rubisco decreased. Across sites and months, the optimal leaf nitrogen content was close to the actual leaf nitrogen content and explained the variation in actual leaf nitrogen by 64 %. Sensitivity analysis showed that the effect of temperature on optimal nitrogen content was similar in magnitude to that of irradiance.

Conclusions

Understorey evergreen species regulate leaf nitrogen content so as to maximize nitrogen-use efficiency in daily carbon gain under changing irradiance and temperature conditions.     

A comparative analysis of leaf shape of wheat, barley and maize using an empirical shape model

Background and Aims

The phenotypes of grasses show differences depending on growth conditions and ontogenetic stage. Understanding these responses and finding suitable mathematical formalizations are an essential part of the development of plant and crop models. Usually, a marked change in architecture between juvenile and adult plants is observed, where dimension and shape of leaves are likely to change. In this paper, the plasticity of leaf shape is analysed according to growth conditions and ontogeny.

Methods

Leaf shape of Triticum aestivum, Hordeum vulgare and Zea mays cultivars grown under varying conditions was measured using digital image processing. An empirical leaf shape model was fitted to measured shape data of single leaves. Obtained values of model parameters were used to analyse the patterns in leaf shape.

Key Results

The model was able to delineate leaf shape of all studied species. The model error was small. Differences in leaf shape between juvenile and adult leaves in T. aestivum and H. vulgare were observed. Varying growth conditions impacted leaf dimensions but did not impact leaf shape of the respective species.

Conclusions

Leaf shape of the studied T. aestivum and H. vulgare cultivars was remarkably stable for a comparable ontogenetic stage (leaf rank), but differed between stages. Along with other aspects of grass architecture, leaf shape changed during the transition from juvenile to adult growth phase. Model-based analysis of leaf shape is a method to investigate these differences. Presented results can be integrated into architectural models of plant development to delineate leaf shape for different species, cultivars and environmental conditions.     

Recovery dynamics of growth, photosynthesis and carbohydrate accumulation after de-submergence: a comparison between two wetland plants showing escape and quiescence strategies

Background and Aims

The capacity for fast-growth recovery after de-submergence is important for establishment of riparian species in a water-level-fluctuation zone. Recovery patterns of two wetland plants, Alternanthera philoxeroides and Hemarthria altissima, showing ‘escape’ and ‘quiescence’ responses, respectively, during submergence were investigated.

Methods

Leaf and root growth and photosynthesis were monitored continuously during 10 d of recovery following 20 d of complete submergence. Above- and below-ground dry weights, as well as carbohydrate concentrations, were measured several times during the experiment.

Key Results

Both species remobilized stored carbohydrate during submergence. Although enhanced internode elongation depleted the carbohydrate storage in A. philoxeroides during submergence, this species resumed leaf growth 3 d after de-submergence concomitant with restoration of the maximal photosynthetic capacity. In contrast, some sucrose was conserved in shoots of H. altissima during submergence, which promoted rapid re-growth of leaves 2 d after de-submergence and earlier than the full recovery of photosynthesis. The recovery of root growth was delayed by 1–2 d compared with leaves in both species.

Conclusions

Submergence tolerance of the escape and quiescence strategies entails not only the corresponding regulation of growth, carbohydrate catabolism and energy metabolism during submergence but also co-ordinated recovery of photosynthesis, growth and carbohydrate partitioning following de-submergence.     

Morphological and molecular identification to secure cultivar maintenance and management of self-sterile Rubus arcticus

Background and Aims

Preservation of cultivar purity creates a particular challenge for plants that are self-incompatible, require insects for cross-pollination, and have easily germinating seeds and vigorously spreading rhizomes. As the fields must be planted with mixed populations, and a balance must be maintained between the cultivars to achieve effective pollination, methods for field monitoring of the relative density of different cultivars must be practical. Furthermore, a DNA-based method is needed for cultivar verification in the collections and outside of the growing season. The aim of this study was to develop both types of methods for Rubus arcticus (arctic bramble).

Methods

Morphological parameters were measured from six cultivars grown on three farms. Observations from the flowers and fruits included: petal and sepal number, flower diameter, arrangement of petals, size of calyx in relation to corolla, fruit weight, yield and soluble sugars. Observations from the leaves included: width and height of middle leaflet, shape of the base of terminal leaflet, shape of terminal leaflet, leaf margin serration and fingertip touch. The applicability of simple sequence repeat (SSR) or microsatellite DNA markers developed for red raspberry was tested on eight arctic bramble cultivars.

Key Results and Conclusions

Morphological and molecular identification methods were developed for R. arcticus. The best morphological characteristics were the length-to-width ratio of the middle leaflet and leaf margin serration. A particular characteristic, fingertip touch, was shown by electron microscopy to be related to the density and quality of the leaf hairs. Red raspberry SSR marker no. 126 proved to be applicable for differentiation of the eight arctic bramble cultivars tested. These identification methods are critical to secure the maintenance and management of R. arcticus. However, the challenges faced and approaches taken are equally applicable to other species with similar biology.     

Silica nanoparticles aid in structural leaf coloration in the Malaysian tropical rainforest understorey herb Mapania caudata

Background and Aims

Blue-green iridescence in the tropical rainforest understorey sedge Mapania caudata creates structural coloration in its leaves through a novel photonic mechanism. Known structures in plants producing iridescent blues consist of altered cellulose layering within cell walls and in special bodies, and thylakoid membranes in specialized plastids. This study was undertaken in order to determine the origin of leaf iridescence in this plant with particular attention to nano-scale components contributing to this coloration.

Methods

Adaxial walls of leaf epidermal cells were characterized using high-pressure-frozen freeze-substituted specimens, which retain their native dimensions during observations using transmission and scanning microscopy, accompanied by energy-dispersive X-ray spectroscopy to identify the role of biogenic silica in wall-based iridescence. Biogenic silica was experimentally removed using aqueous Na₂CO₃ and optical properties were compared using spectral reflectance.

Key Results and Conclusions

Blue iridescence is produced in the adaxial epidermal cell wall, which contains helicoid lamellae. The blue iridescence from cell surfaces is left-circularly polarized. The position of the silica granules is entrained by the helicoid microfibrillar layers, and granules accumulate at a uniform position within the helicoids, contributing to the structure that produces the blue iridescence, as part of the unit cell responsible for 2 ° Bragg scatter. Removal of silica from the walls eliminated the blue colour. Addition of silica nanoparticles on existing cellulosic lamellae is a novel mechanism for adding structural colour in organisms.     

Nitrogen stress response of a hybrid species: a gene expression study

Background and Aims

Low soil fertility limits growth and productivity in many natural and agricultural systems, where the ability to sense and respond to nutrient limitation is important for success. Helianthus anomalus is an annual sunflower of hybrid origin that is adapted to desert sand-dune substrates with lower fertility than its parental species, H. annuus and H. petiolaris. Previous studies have shown that H. anomalus has traits generally associated with adaptation to low-fertility habitats, including a lower inherent relative growth rate and longer leaf lifetime.

Methods

Here, a cDNA microarray is used to identify gene expression differences that potentially contribute to increased tolerance of low fertility of the hybrid species by comparing the nitrogen stress response of all three species with high- and low-nutrient treatments.

Key Results

Relative to the set of genes on the microarray, the genes showing differential expression in the hybrid species compared with its parents are enriched in stress-response genes, developmental genes, and genes involved in responses to biotic or abiotic stimuli. After a correction for multiple comparisons, five unique genes show a significantly different response to nitrogen limitation in H. anomalus compared with H. petiolaris and H. annuus. The Arabidopsis thaliana homologue of one of the five genes, catalase 1, has been shown to affect the timing of leaf senescence, and thus leaf lifespan.

Conclusions

The five genes identified in this analysis will be examined further as candidate genes for the adaptive stress response in H. anomalus. Genes that improve growth and productivity under nutrient stress could be used to improve crops for lower soil fertility which is common in marginal agricultural settings.     

Leaf trait co-ordination in relation to construction cost, carbon gain and resource-use efficiency in exotic invasive and native woody vine species

Background and Aims

Success of invasive plant species is thought to be linked with their higher leaf carbon fixation strategy, enabling them to capture and utilize resources better than native species, and thus pre-empt and maintain space. However, these traits are not well-defined for invasive woody vines.

Methods

In a glass house setting, experiments were conducted to examine how leaf carbon gain strategies differ between non-indigenous invasive and native woody vines of south-eastern Australia, by investigating their biomass gain, leaf structural, nutrient and physiological traits under changing light and moisture regimes.

Key Results

Leaf construction cost (CC), calorific value and carbon : nitrogen (C : N) ratio were lower in the invasive group, while ash content, N, maximum photosynthesis, light-use efficiency, photosynthetic energy-use efficiency (PEUE) and specific leaf area (SLA) were higher in this group relative to the native group. Trait plasticity, relative growth rate (RGR), photosynthetic nitrogen-use efficiency and water-use efficiency did not differ significantly between the groups. However, across light resource, regression analyses indicated that at a common (same) leaf CC and PEUE, a higher biomass RGR resulted for the invasive group; also at a common SLA, a lower CC but higher N resulted for the invasive group. Overall, trait co-ordination (using pair-wise correlation analyses) was better in the invasive group. Ordination using 16 leaf traits indicated that the major axis of invasive-native dichotomy is primarily driven by SLA and CC (including its components and/or derivative of PEUE) and was significantly linked with RGR.

Conclusions

These results demonstrated that while not all measures of leaf resource traits may differ between the two groups, the higher level of trait correlation and higher revenue returned (RGR) per unit of major resource need (CC) and use (PEUE) in the invasive group is in line with their rapid spread where introduced.     

Plasticity of stomatal distribution pattern and stem tracheid dimensions in Podocarpus lambertii: an ecological study

Background and Aims

Leaf and wood plasticity are key elements in the survival of widely distributed plant species. Little is known, however, about variation in stomatal distribution in the leaf epidermis and its correlation with the dimensions of conducting cells in wood. This study aimed at testing the hypothesis that Podocarpus lambertii, a conifer tree, possesses a well-defined pattern of stomatal distribution, and that this pattern can vary together with the dimensions of stem tracheids as a possible strategy to survive in climatically different sites.

Methods

Leaves and wood were sampled from trees growing in a cold, wet site in south-eastern Brazil and in a warm, dry site in north-eastern Brazil. Stomata were thoroughly mapped in leaves from each study site to determine a spatial sampling strategy. Stomatal density, stomatal index and guard cell length were then sampled in three regions of the leaf: near the midrib, near the leaf margin and in between the two. This sampling strategy was used to test for a pattern and its possible variation between study sites. Wood and stomata data were analysed together via principal component analysis.

Key Results

The following distribution pattern was found in the south-eastern leaves: the stomatal index was up to 25 % higher in the central leaf region, between the midrib and the leaf margin, than in the adjacent regions. The inverse pattern was found in the north-eastern leaves, in which the stomatal index was 10 % higher near the midrib and the leaf margin. This change in pattern was accompanied by smaller tracheid lumen diameter and length.

Conclusions

Podocarpus lambertii individuals in sites with higher temperature and lower water availability jointly regulate stomatal distribution in leaves and tracheid dimensions in wood. The observed stomatal distribution pattern and variation appear to be closely related to the placement of conducting tissue in the mesophyll.     

Juvenility and flowering of Brunonia australis (Goodeniaceae) and Calandrinia sp. (Portulacaceae) in relation to vernalization and daylength

Background and Aims

The time at which plants are transferred to floral inductive conditions affects the onset of flowering and plant morphology, due to juvenility. Plants of Brunonia australis and Calandrinia sp. were used to investigate whether Australian native ephemeral species show a distinct juvenile phase that can be extended to increase vegetative growth and flowering.

Methods

The juvenile phase was quantified by transferring seedlings from less inductive (short day and 30/20°C) to inductive (vernalization or long day) conditions at six different plant ages ranging from 4 to 35 d after seed germination. An increase in days to first visible floral bud and leaf number were used to signify the end of juvenility.

Key Results

Brunonia australis was receptive to floral inductive long day conditions about 18–22 d after seed germination, whereas plants aged 4–35 d appeared vernalization sensitive. Overall, transferring plants of B. australis from short to long day conditions reduced the time to anthesis compared with vernalization or constant short day conditions. Calandrinia sp. showed a facultative requirement for vernalization and an insensitive phase was not detected. Floral bud and branch production increased favourably as plant age at time of transfer to inductive conditions increased. Younger plants showed the shortest crop production time.

Conclusions

Both species can perceive the vernalization floral stimulus from a very young age, whereas the photoperiodic stimulus is perceived by B. australis after a period of vegetative growth. However, extending the juvenile phase can promote foliage development and enhance flower production of both species.     

Effects of mating system on adaptive potential for leaf morphology in Crepis tectorum (Asteraceae)

Background and Aims

A shift from outcrossing to selfing is thought to reduce the long-term survival of populations by decreasing the genetic variation necessary for adaptation to novel ecological conditions. However, theory also predicts an increase in adaptive potential as more of the existing variation becomes expressed as homozygous genotypes. So far, relatively few studies have examined how a transition to selfing simultaneously affects means, variances and covariances for characters that might be under stabilizing selection for a spatially varying optimum, e.g. characters describing leaf morphology.

Methods

Experimental crosses within an initially self-sterile population of Crepis tectorum were performed to produce an outbred and inbred progeny population to assess how a shift to selfing affects the adaptive potential for measures of leaf morphology, with special emphasis on the degree of leaf dissection, a major target of diversifying selection within the study species.

Key Results

Three consecutive generations of selfing had a minor impact on survival, the total number of heads produced and the mean leaf phenotype, but caused a proportional increase in the genetic (co)variance matrix for foliar characters. For the degree of leaf dissection, the lowest 50th percentile of the inbred progeny population showed a disproportionate increase in the genetic variance, consistent with the recessive nature of the weakly lobed phenotype observed in interpopulation crosses. Comparison of inbreeding response with large-scale patterns of variation indicates a potential for selection in a (recently) inbred population to drive a large evolutionary reduction in degree of leaf dissection by increasing the frequency of particular sibling lines.

Conclusions

The results point to a positive role for inbreeding in phenotypic evolution, at least during or immediately after a rapid shift in mating system.     

Foliage nitrogen turnover: differences among nitrogen absorbed at different times by Quercus serrata saplings

Background and Aims

Nitrogen turnover within plants has been intensively studied to better understand nitrogen use strategies. However, differences among the nitrogen absorbed at different times are not completely understood and the fate of nitrogen absorbed during winter is largely uncharacterized. In the present study, nitrogen absorbed at different times of the year (growing season, winter and previous growing season) was traced, and the within-leaf nitrogen turnover of a temperate deciduous oak Quercus serrata was investigated.

Methods

The contributions of nitrogen absorbed at the three different times to leaf construction, translocation during the growing season, and the leaf-level resorption efficiency during leaf senescence were compared using ¹⁵N.

Key Results

Winter- and previous growing season-absorbed nitrogen significantly contributed to leaf construction, although the contribution was smaller than that of growing season-absorbed nitrogen. On the other hand, the leaf-level resorption efficiency of winter- and previous growing season-absorbed nitrogen was higher than that of growing season-absorbed nitrogen, suggesting that older nitrogen is better retained in leaves than recently absorbed nitrogen.

Conclusions

The results demonstrate that nitrogen turnover in leaves varies with nitrogen absorption times. These findings are important for understanding plant nitrogen use strategies and nitrogen cycles in forest ecosystems.