Leaf Life Span Research Articles

葉寿命研究の歴史と近況(企画趣旨, なぜいま葉寿命なのか?)

葉寿命研究の歴史と近況(企画趣旨, なぜいま葉寿命なのか?)

Cost-benefit modelを用いた最適葉寿命モデル : 最適戦略基準の検討( なぜいま葉寿命なのか?)

Cost-benefit modelを用いた最適葉寿命モデル : 最適戦略基準の検討( なぜいま葉寿命なのか?)

Plant traits and environment: floating leaf blade production and turnover of waterlilies.

Floating leaf blades of waterlilies fulfill several functions in wetland ecosystems by production, decomposition and turnover as well as exchange processes. Production and turnover rates of floating leaf blades of three waterlily species, Nuphar lutea (L.) Sm., Nymphaea alba L. and Nymphaea candida Presl, were studied in three freshwater bodies, differing in trophic status, pH and alkalinity. Length and percentages of leaf loss of marked leaf blades were measured weekly during the growing season. Area and biomass were calculated based on leaf length and were used to calculate the turnover rate of floating leaf blades. Seasonal changes in floating leaf production showed that values decreased in the order: Nymphaea alba, Nuphar lutea, Nymphaea candida. The highest production was reached for Nuphar lutea and Nymphaea alba in alkaline, eutrophic water bodies. The production per leaf was relatively high for both species in the acid water body. Nymphaea candida showed a very short vegetation period and low turnover rates. The ratio Total potential leaf biomass/Maximum potential leaf biomass (P/Bmax) of the three species ranged from 1.35–2.25. The ratio Vegetation period (Period with floating leaves)/Mean leaf life span ranged from 2.94–4.63, the ratio Growth period (Period with appearance of new floating leaves)/Vegetation period from 0.53–0.73. The clear differences between Nymphaea candida versus Nuphar lutea and Nymphaea alba, may be due to adaptations of Nymphaea candida to an Euro-Siberic climate with short-lasting summer conditions.

Long-term simulated nitrogen deposition alters the plant cover dynamics of a Mediterranean rosemary shrubland in Central Spain through defoliation.

Nitrogen (N) deposition due to anthropogenic pollution is a major driver of the global biodiversity loss. We studied the effect of experimental N and phosphorus (P) fertilization (0, 10, 20, and 50kg N ha-1 year-1 and 14kg P ha-1year-1 over the background deposition levels) on plant cover dynamics of a rosemary (Rosmarinus officinalis L.) shrubland after 8years of nutrient addition in a semiarid Mediterranean ecosystem from Central Spain. We specifically aimed at testing whether N deposition has the potential to influence the observed expanding trend of woody vegetation into areas dominated by grassland, biological soil crusts, and bare soil. Our results show that N addition loads above 10kg N ha-1 year-1 reverted the cover dynamics of shrubs. Under N addition conditions, N was no longer a limiting nutrient and other elements, especially P and calcium, determined the seasonal growth of young twigs. Interestingly, N fertilization did not inhibit the growth of young shoots; our estimates point to a reduced rosemary leaf lifespan that is driving individuals to death. This may be triggered by long-term accumulation of N compounds in leaves, suggesting the need to consider the old organs and tissues in long-lived perennial plants, where N toxicity effects could be more mediated by accumulation processes. Shrublands are a widely distributed ecosystem type in biodiverse Mediterranean landscapes, where shrubs play a key role as nurse plants. Therefore, the disappearance of shrublands may accelerate the biodiversity loss associated with other global change drivers, hamper the recruitment of seedlings of woody species, and, as a consequence, accelerate desertification.

Early life conditions and precipitation influence the performance of widespread understorey herbs in variable light environments

Summary The understorey of tropical forests is heterogeneous across time, and plants that inhabit this layer may exhibit adaptations (e.g. trait plasticity) that enable them to exploit this variability to their advantage. We tested the hypothesis that two widespread understorey herbs would perform equally well in a variable as in a constant environment, using a 2‐year shade‐house experiment. We measured demographic traits (growth and survival), a physiological trait (maximum photosynthetic capacity), and life‐history traits (leaf life span and biomass allocation) of Heliconia tortuosa and Calathea crotalifera. We investigated how these traits were affected by light availability at the seedling stage, precipitation, and whether individuals experienced a constant or variable light environment. Whether or not a variable environment was favourable for plants depended upon precipitation and the environment in which individuals started life. At low precipitation, plants in a variable light environment grew more than those in a constant environment, but only when individuals had lived as seedlings in low light. At high precipitation, plants in a constant environment grew more than those in a variable environment, regardless of early conditions. Survival was lower in a variable environment at low precipitation, and more so at high precipitation. Photosynthetic capacity was lower for individuals in a variable environment than in a constant environment when they had lived in high light as seedlings. Calathea grew faster and survived more poorly than Heliconia, independently of the treatments. Calathea grew more at high than low precipitation while Heliconia grew more at low than high precipitation. Leaf life span and biomass allocation did not differ among treatments, although Calathea had a significantly greater proportion of its biomass above‐ground vs. that of Heliconia. Synthesis. Environmental variability had a neutral or positive effect on biomass allocation, photosynthetic capacity, and leaf life span for these species. Survival was the only trait that was always lower in a variable environment. The effect of environmental variability was dependent on early life conditions as well as precipitation, suggesting that generalist species may experience high fitness as forest environments become more variable by maintaining high growth at the expense of survival.

Physiological and structural tradeoffs underlying the leaf economics spectrum.

The leaf economics spectrum (LES) represents a suite of intercorrelated leaf traits concerning construction costs per unit leaf area, nutrient concentrations, and rates of carbon fixation and tissue turnover. Although broad trade-offs among leaf structural and physiological traits have been demonstrated, we still do not have a comprehensive view of the fundamental constraints underlying the LES trade-offs. Here, we investigated physiological and structural mechanisms underpinning the LES by analysing a novel data compilation incorporating rarely considered traits such as the dry mass fraction in cell walls, nitrogen allocation, mesophyll CO2 diffusion and associated anatomical traits for hundreds of species covering major growth forms. The analysis demonstrates that cell wall constituents are major components of leaf dry mass (18-70%), especially in leaves with high leaf mass per unit area (LMA) and long lifespan. A greater fraction of leaf mass in cell walls is typically associated with a lower fraction of leaf nitrogen (N) invested in photosynthetic proteins; and lower within-leaf CO2 diffusion rates, as a result of thicker mesophyll cell walls. The costs associated with greater investments in cell walls underpin the LES: long leaf lifespans are achieved via higher LMA and in turn by higher cell wall mass fraction, but this inevitably reduces the efficiency of photosynthesis.

Morphogenic, structural, and productive characteristics of Cenchrus ciliaris L. grown using different levels of caprine and ovine biofertilize

The aim of this study was to evaluate the morphogenic, structural, and productive characteristics of Cenchrus ciliaris L. in response to different doses of biofertilizers that originate from the anaerobic biodigestion of goat and sheep fecal matter on Yellow Argisoil. The experiment was performed in a greenhouse, in a completely randomized factorial design using two types of biofertilizers and five dosages (0%, 2.5%, 5.0%, 7.5%, and 10% of soil volume), with four repeats, for the duration of two vegetative cycles. In the first cycle, there was an increase in leaf lifespan and leaf/stem ratio with the use of 10% and 7.5% caprine biofertilizer, respectively. From the second cycle, doses of biofertilizers between 7.5% and 10% led to an improvement in the development of the morphogenic variables - leaf sprouting rate, leaf and pseudostem elongation rate, leaf lifespan, and senescence, as well as the structural characteristics - expanded leaf length and width, number of sprouting, expanded and live leaves, and expanded/expanding leaf ratio, resulting in a 73% increase in biomass. It was concluded that the use of biofertilizers improves the main morphogenic and structural characteristics of Cenchrus ciliaris L., resulting in increased fodder mass yield.

High-Throughput and Computational Study of Leaf Senescence through a Phenomic Approach.

Leaf senescence is influenced by its life history, comprising a series of developmental and physiological experiences. Exploration of the biological principles underlying leaf lifespan and senescence requires a schema to trace leaf phenotypes, based on the interaction of genetic and environmental factors. We developed a new approach and concept that will facilitate systemic biological understanding of leaf lifespan and senescence, utilizing the phenome high-throughput investigator (PHI) with a single-leaf-basis phenotyping platform. Our pilot tests showed empirical evidence for the feasibility of PHI for quantitative measurement of leaf senescence responses and improved performance in order to dissect the progression of senescence triggered by different senescence-inducing factors as well as genetic mutations. Such an establishment enables new perspectives to be proposed, which will be challenged for enhancing our fundamental understanding on the complex process of leaf senescence. We further envision that integration of phenomic data with other multi-omics data obtained from transcriptomic, proteomic, and metabolic studies will enable us to address the underlying principles of senescence, passing through different layers of information from molecule to organism.

Functional leaf traits and leaf economics in the Paleogene — A case study for Central Europe

Numerous data based on extant vegetation reveal global patterns of relationships between functional leaf traits and climate. Leaf life span (LLS), i.e. evergreen vs. deciduous leaves, represents a central parameter linking functional traits related to the global leaf economics spectrum. Paleogene climate transitions are therefore expected to be reflected by functional leaf traits and leaf economics. In this study, fossil floras from six sites in Central Europe dating back to the Paleocene, late Eocene, Early and Late Oligocene are studied, addressing the following questions: 1) How does leaf economics and LLS change through the Paleogene? 2) How do various functional leaf traits change through the Paleogene, and how do they relate to leaf economics? 3) Are changes in leaf functional traits consistent with climate reconstructions from proxy data? As a proxy for LLS change, leaf mass per area (LMA) was determined indirectly. The results show the late Eocene site to stand out from all other sites, by tending towards lower leaf size/width, higher leaf mass per area (LMA), a tendency towards untoothed leaf margins, a high frequency of looped secondary veins, an almost complete absence of lobed leaves, and a low proportion of toothed pinnate leaves with non-looped secondaries. The LMA peak at the Eocene site is caused by untoothed leaf types tending to higher LMA and by an increase in LMA of toothed leaf types, compared to the Paleocene and Oligocene. The proportion of lobed leaves, which usually tend to lower LMA and therefore to deciduousness, increases from the older to the younger Oligocene sites. The results demonstrate that LMA data are valuable for evaluating functional/ecological contexts of architectural traits of fossil leaves. The results are consistent with a post-Eocene cooling trend with mild winters and the development of a drier season, both favoring a trend towards deciduous vegetation.

<b>Efficiency of the use of ryegrass by heifers in response to the receival of supplement

The intensity and frequency of defoliation of ryegrass (Lolium multiflorum Lam.) by beef heifers exclusively grazing or receiving corn grain or extruded commercial product with fat as supplement were evaluated. The grazing method was put-and-take stocking. The experimental design was completely randomized following a repeated measure arrangement, three treatments and two area replications. The phyllochron of ryegrass was 140.1 and 303.4°GD the leaf lifespan. The intensity (67.0 %) and the frequency of defoliation (6.4 days) were similar, regardless of supply or no of energetic supplement for grazing heifers. The defoliation frequency indicates that the heifers can return about 3.3 times to the same tiller during the leaf lifespan. Considering the different categories of leaf blades, the supply of extruded fat causes the heifers to remove higher proportion (66.5 %) of expanding leaf blades. On ryegrass pasture, the efficiency of forage use is not modified when heifers receive energetic supplement.

Inconsistent intraspecific pattern in leaf life span along nitrogen-supply gradient.

Leaf life span (LLS) has long been hypothesized to plastically increase with decreasing nitrogen (N) supply from soil to maximize N retention, carbon assimilation, and fitness; however, accumulating evidence shows no consistent trend. The apparent inconsistencies are explained by a recent model that assumes LLS has a hump-shaped quadratic response to the N-supply gradient. The available evidence mostly originates from comparisons of LLS at only two levels of N availability, and the hypothesis remains unanswered. We investigated LLS of two asteraceous forbs (Adenocaulon himalaicum and Xanthium canadense) experimentally grown at eight levels of N supply, which covered a range of N supply in their natural habitats. We additionally conducted a literature search to retrieve studies reporting LLS response along an N-supply gradient. The LLS of neither species showed a hump-shaped response along the N-supply gradient. Past studies examining the LLS of an aquatic forb and terrestrial shrubs and trees along the N-supply gradient (more than four levels of N supply) also refuted the hypothesis. The LLS of a single species exhibited neither an increase nor a hump-shaped response to decreased N supply in a variety of life forms. Comparisons at only a few N levels are misleading with regard to LLS response to N supply.

Leaf Litterfall and Decomposition ofPolylepis reticulatain the Treeline of the Ecuadorian Andes

Leaf litterfall contributes significantly to carbon fluxes in forests. A crucial open question for the sustainability of mountain forests is how climate change will affect this and other carbon fluxes (eg photosynthesis and respiration). Leaf litterfall and decomposition of Polylepis reticulata, an endemic species of the Andes, were analyzed during a period of 1 year at 6 experimental plots located in the Andean paramo between 3700 and 3900 m above sea level in Cajas National Park, Ecuador. Litterfall was collected in each plot using 5 randomly distributed traps. Every trap had a 40-cm diameter (0.125 m2) and was suspended 0.8 to 1.0 m above the ground. The decomposition rate of the leaf litter was analyzed using litter bags. Eighteen bags with approximately 20 g of dry litter were placed in the litter layer in each experimental plot and collected 30, 60, 90, 150, 210, 300, and 365 days after they were installed. The mean annual litterfall recorded was 3.77 Mg ha−1, representing 51% of the leaf biomass present in the canopy, so the leaf life span of P. reticulata in Cajas National Park is 1.98 years. Litterfall occurred all year, with no significant seasonal pattern. The mean decomposition rate (k) obtained for this study period was 0.38 year−1. This study contributes to the information gap on litterfall and decomposition in natural forests located at the highest elevations in the world.

The association of leaf lifespan and background insect herbivory at the interspecific level.

Herbivory is well known to be a major selective pressure that affects plant communities, but the leaf traits that mediate variations in herbivory at the interspecific level remain controversial. We collected published data on background insect herbivory and leaf traits from a wide variety of species to test the hypothesis that species with intermediate leaf lifespans, lower fiber, and higher nutrient contents in leaves should have higher levels of herbivory. We found that at the interspecific level herbivory had a hump-shaped relationship with leaf lifespan and a positive relationship with leaf size. Surprisingly, our data show that nutritional traits have little relationship to herbivory. Our study provides new insights relevant to the recent debate on leaf trait-herbivory relationships. These findings are especially helpful in explaining the general patterns of herbivory detected on the global scale.

Leaf morphogenesis influences nutritive-value dynamics of tall fescue (Lolium arundinaceum) cultivars of different leaf softness

The objective of this study was to compare the dynamics of neutral detergent fibre (NDF), and the 24-h in-vitro digestibility of NDF (NDFD) and dry matter (DMD) in leaf blades of two tall fescue (Lolium arundinaceum (Schreb.) Darbysh.) cultivars of different leaf softness: a soft- and a tough-leaved cultivar. The experiment was conducted during the summer regrowth of three replicated, dense mini-swards per cultivar arranged in a completely randomised design, all grown under non-limiting water, nitrogen and phosphorus. Cultivars were harvested eight times over 14 weeks to measure morphogenetic traits and nutritive value in six predefined leaf-age categories (from growing to complete senescence). The leaf lifespan and leaf length of the first three successive leaves were measured on 30 marked tillers throughout the experiment. Following analysis of variance, linear regression models were fitted to describe variations of NDF, NDFD and DMD with increasing leaf age and leaf length. Similar leaf NDF contents were found for the two cultivars, which remained stable throughout the leaf lifespan and increased markedly during leaf senescence. Leaf NDFD and leaf DMD both declined with increasing leaf age and length for the two cultivars. However, owing to shorter leaf lifespan of the soft-leaved cultivar, this decline in leaf NDFD and leaf DMD was faster for the soft- than for the tough-leaved cultivar. These results suggest that the soft-leaved cultivar will require more frequent defoliations than the tough-leafed cultivar to prevent decreases in nutritive value.

Community aggregated traits disclose functional responses to seasonal resource fluctuations and spatial heterogeneity

Aim To investigate changes in community-level functional responses to rainfall seasonality in Neotropical grasslands through the analysis of community aggregated traits. Location Semi-natural grasslands in the Rio Grande de Matagalpa watershed, Nicaragua. Methods We measured 14 functional response traits that are indicators of plant resource-use strategies, across 32 herbaceous and four woody species in eight permanent plots: leaf size, specific leaf area, leaf dry matter content, leaf lifespan, foliar concentrations of P, N, Ca, K and Mg, plant height, lateral spread by clonal growth, root depth, start of flowering period and length of the period from flowering to seed shed. We calculated the community-weighted means (CWM) with trait values weighted by species cover for five different paddocks 11 times (between 2 Jul 2007 and 26 May 2008), and grouped them into four periods according to rainfall seasonality: early rainy season, late rainy season, early dry season and late dry season. Results Community aggregated values of specific leaf area, leaf dry matter content, leaf lifespan, foliar concentrations of P, N, Ca, K and Mg, length of the period from flowering to seed shed, and to a lesser extent lateral spread by clonal growth responded to temporal variations in rainfall, and corresponded to a considerable extent to expectations based on plant resource-use strategies along resource supply gradients. Community aggregated values of specific leaf area and foliar nutrient concentration were higher in the rainy season than in the dry season, while we observed an opposite trend with leaf dry matter content and leaf lifespan. Conclusions Semi-natural grasslands of central Nicaragua experience shifts in plant trait dominance that correspond with changes in resource supply given by rainfall seasonality, indicating that the stabilizing function of net primary productivity found in earlier studies can in part be a consequence of temporal differentiation in functional responses.

Predicting vegetation type through physiological and environmental interactions with leaf traits: evergreen and deciduous forests in an earth system modeling framework.

Earth system models are incorporating plant trait diversity into their land components to better predict vegetation dynamics in a changing climate. However, extant plant trait distributions will not allow extrapolations to novel community assemblages in future climates, which will require a mechanistic understanding of the trade-offs that determine trait diversity. In this study, we show how physiological trade-offs involving leaf mass per unit area (LMA), leaf lifespan, leaf nitrogen, and leaf respiration may explain the distribution patterns of evergreen and deciduous trees in the temperate and boreal zones based on (1) an evolutionary analysis of a simple mathematical model and (2) simulation experiments of an individual-based dynamic vegetation model (i.e., LM3-PPA). The evolutionary analysis shows that these leaf traits set up a trade-off between carbon- and nitrogen-use efficiency at the scale of individual trees and therefore determine competitively dominant leaf strategies. As soil nitrogen availability increases, the dominant leaf strategy switches from one that is high in nitrogen-use efficiency to one that is high in carbon-use efficiency or, equivalently, from high-LMA/long-lived leaves (i.e., evergreen) to low-LMA/short-lived leaves (i.e., deciduous). In a region of intermediate soil nitrogen availability, the dominant leaf strategy may be either deciduous or evergreen depending on the initial conditions of plant trait abundance (i.e., founder controlled) due to feedbacks of leaf traits on soil nitrogen mineralization through litter quality. Simulated successional patterns by LM3-PPA from the leaf physiological trade-offs are consistent with observed successional dynamics of evergreen and deciduous forests at three sites spanning the temperate to boreal zones.

Incorporation of plant traits in a land surface model helps explain the global biogeographical distribution of major forest functional types

AbstractAimsContrasting leaf types with different leaf life spans represent different adaptive strategies in plants. Previous studies explained the adaptive advantages of different strategies on the basis of environmental climatic limits, but could not account for the observed co‐dominance of multiple plant functional types (PFTs) in many regions. Here we used a process‐based model to explore whether observed inter‐ and intra‐PFT variation in plant traits can explain global biogeographical variation in the dominance and co‐dominance of major forest types.LocationWorld‐wide.MethodsWe identified four important plant traits: leaf N concentration, leaf life span, fraction of net primary production (NPP) allocated to leaves and plant basal respiration rate. We incorporated means and variances of these plant traits from trait databases into the Community Atmosphere–Biosphere–Land Exchange model. We then predicted the dominant PFT or PFT mixture for global forested grid cells, using NPP as a proxy for growth rate, and considering three PFTs: evergreen needleleaf forests (ENF), evergreen broadleaf forests (EBF) and deciduous broadleaf forests (DBF). We compared results with: (1) those from simulations that did not account for trait variance; (2) results from an empirical model based purely on mean annual temperature; and (3) data from remotely sensed observations.ResultsOur estimates of the fractions of land area covered by major forest types were consistent with observation; i.e. ENFs dominate in boreal regions, EBFs dominate in tropical regions and DBFs are distributed widely across a broad range of environmental conditions. We also showed that co‐dominance of different forest PFTs cannot be reproduced without considering variation in plant traits.Main conclusionsGlobal trait data are useful for representing underlying plant strategies and functional diversity. Variation in key plant traits explains significant fractions of global biogeographical variation of three major forest types. Future developments in dynamic global vegetation modelling will benefit from the inclusion of plant trait variation.

<b>Morphogenesis and structure of Tifton 85 cultivated in subtropical climate and fertilized with nitrogen

The experiment evaluated the morphogenesis and structure of Tifton 85 (Cynodon spp.) cultivated in subtropical climate and fertilized with nitrogen (N). The experiment was a completely randomized design with four levels of N (Zero; 75; 150 or 225 kg ha-1) in nine replicates per area. The experimental animals were Suffolk female lambs. The grazing method was continuous to maintain the sward height at 15 cm ± 10%. The stem expansion increased by 0.000027 cm degree-days-1 to each kg N applied (linear model). According to nonlinear model, the highest stem expansion (0.0226 cm degree-days-1) was observed with the use 220.1 kg ha-1 N. The leaf lifespan fitted a linear regression model, with increase of 50 degree-days leaf -1, comparing the levels zero and 225 kg ha-1 of N. According to nonlinear regression model, the longest leaf lifespan (407.1 degree-days) was observed with the use 208.8 kg ha-1 N. With the maintenance of sward height at approximately 15 cm, we recommended to use 200 kg ha-1 N in Tifton 85 cultivated in subtropical climate.

Leaf traits of dipterocarp species with contrasting distributions across a gradient of nutrient and light availability

Background: Tree species composition at the landscape scale is often tightly associated with underlying soil type in tropical forests. Changes in soil type may have effects on forest structure that drive changes in both light and soil resource availability, since light availability in the understorey tends to be lower in more fertile sites. Plant functional traits may determine species distributions across gradients of light and soil resource availability.Aims: To test whether tree species with contrasting distributions exhibit leaf traits that reflect adaptation to the resources most limiting in their native environment.Methods: We measured foliar nutrient concentrations, stomatal density, leaf δ13C values, leaf mass per area, and leaf lifespan for saplings of nine common dipterocarp species at Sepilok Forest Reserve, Malaysian Borneo, possessing varying associations to soil resource habitats.Results: Species specialised in their adult distribution to nutrient-poor sandstone soils had traits indicative of a nutrient conservation strategy. Species specialised to more fertile alluvial soils had a wider spectrum of leaf N and P concentrations and LL, reflecting greater variance in strategies for resource acquisition and use among species in this habitat.Conclusions: Understorey light regimes co-vary with soil type, and both light and soil resource availability influence leaf trait adaptations that may contribute to species–habitat associations.

Apparent Overinvestment in Leaf Venation Relaxes Leaf Morphological Constraints on Photosynthesis in Arid Habitats.

Leaf veins supply the mesophyll with water that evaporates when stomata are open to allow CO2 uptake for photosynthesis. Theoretical analyses suggest that water is optimally distributed in the mesophyll when the lateral distance between veins (dx) is equal to the distance from these veins to the epidermis (dy), expressed as dx:dy ≈ 1. Although this theory is supported by observations of many derived angiosperms, we hypothesize that plants in arid environments may reduce dx:dy below unity owing to climate-specific functional adaptations of increased leaf thickness and increased vein density. To test our hypothesis, we assembled leaf hydraulic, morphological, and photosynthetic traits of 68 species from the Eucalyptus and Corymbia genera (termed eucalypts) along an aridity gradient in southwestern Australia. We inferred the potential gas-exchange advantage of reducing dx beyond dy using a model that links leaf morphology and hydraulics to photosynthesis. Our observations reveal that eucalypts in arid environments have thick amphistomatous leaves with high vein densities, resulting in dx:dy ratios that range from 1.6 to 0.15 along the aridity gradient. Our model suggests that, as leaves become thicker, the effect of reducing dx beyond dy is to offset the reduction in leaf gas exchange that would result from maintaining dx:dy at unity. This apparent overinvestment in leaf venation may be explained from the selective pressure of aridity, under which traits associated with long leaf life span, high hydraulic and thermal capacitances, and high potential rates of leaf water transport confer a competitive advantage.