Complementary resource use by tree species in a rain forest tree plantation

Mixed-species tree plantations, composed of high-value native rain forest timbers, are potential forestry systems for the subtropics and tropics that can provide ecological and production benefits. Choices of rain forest tree species for mixtures are generally based on the concept that assemblages of fast-growing and light-demanding species are less productive than assemblages of species with different shade tolerances. We examined the hypothesis that mixtures of two fast-growing species compete for resources, while mixtures of shade-tolerant and shade-intolerant species are complementary. Ecophysiological characteristics of young trees were determined and analyzed with a physiology-based canopy model (MAESTRA) to test species interactions. Contrary to predictions, there was evidence for complementary interactions between two fast-growing species with respect to nutrient uptake, nutrient use efficiency, and nutrient cycling. Fast-growing Elaeocarpus angustifolius had maximum demand for soil nutrients in summer, the most efficient internal recycling of N, and low P use efficiency at the leaf and whole-plant level and produced a large amount of nutrient-rich litter. In contrast, fast-growing Grevillea robusta had maximum demand for soil nutrients in spring and highest leaf nutrient use efficiency for N and P and produced low-nutrient litter. Thus, mixtures of fast-growing G. robusta and E. angustifolius or G. robusta and slow-growing, shade-tolerant Castanospermum australe may have similar or even greater productivity than monocultures, as light requirement is just one of several factors affecting performance of mixed-species plantations. We conclude that the knowledge gained here will be useful for designing large-scale experimental mixtures and commercial forestry systems in subtropical Australia and elsewhere

Evolution of leaf-form in land plants linked to atmospheric CO2 decline in the Late Palaeozoic era

The widespread appearance of megaphyll leaves, with their branched veins and planate form, did not occur until the close of the Devonian period at about 360 Myr ago. This happened about 40 Myr after simple leafless vascular plants first colonized the land in the Late Silurian/Early Devonian, but the reason for the slow emergence of this common feature of present-day plants is presently unresolved. Here we show, in a series of quantitative analyses using fossil leaf characters and biophysical principles, that the delay was causally linked with a 90% drop in atmospheric pCO2 during the Late Palaeozoic era. In contrast to simulations for a typical Early Devonian land plant, possessing few stomata on leafless stems, those for a planate leaf with the same stomatal characteristics indicate that it would have suffered lethal overheating, because of greater interception of solar energy and low transpiration. When planate leaves first appeared in the Late Devonian and subsequently diversified in the Carboniferous period, they possessed substantially higher stomatal densities. This observation is consistent with the effects of the pCO2 on stomatal development and suggests that the evolution of planate leaves could only have occurred after an increase in stomatal density, allowing higher transpiration rates that were sufficient to maintain cool and viable leaf temperatures

Correlation between carbon isotope discrimination and transpiration efficiency in lines of the C4 species Sorghum bicolor in the glasshouse and the field

Transpiration efficiency, W, the ratio of plant carbon produced to water transpired and carbon isotope discrimination of leaf dry matter, Δd, were measured together on 30 lines of the C4 species, Sorghum bicolor, in the glasshouse and on eight lines grown in the field. In the glasshouse, the mean W observed was 4.9 mmol C mol-1 H2O and the range was 0.8 mmol C mol -1 H2O. The mean Δd was 3.0 and the observed range was 0.4‰. In the field, the mean W was lower at 2.8 mmol C mol-1 H2O and the mean Δd was 4.6‰. Significant positive correlations between W and Δd were observed for plants grown in the glasshouse and in the field. The observed correlations were consistent with theory, opposite to those for C3 species, and showed that variation in Δd was an integrated measure of long-term variation in the ratio of intercellular to ambient CO2 partial pressure, pi/pa. Detailed gas exchange measurements of carbon isotope discrimination during CO2 uptake, ΔA, and pi/pa were made on leaves of eight S. bicolorlines. The observed relationship between ΔA and pi/pa was linear with a negative slope of 3.7‰ in ΔA for a unit change in pi/pa. The slope of this linear relationship between ΔA and pi/pa in C4 species is dependent on the leakiness of the CO2 concentrating mechanism of the C4 pathway. We estimated the leakiness (defined as the fraction of CO2 released in the bundle sheath by C4 acid decarboxylations, which is lost by leakage) to be 0.2. We conclude that, although variation in Δd observed in the 30 lines of S. bicolor is smaller than that commonly observed in C3 species, it also reflects variation in transpiration efficiency, W. Among the eight lines examined in detail and in the environments used, there was considerable genotype × environment interaction

Temperature responses of Rubisco from Paniceae grasses provide opportunities for improving C3 photosynthesis.

Enhancing the catalytic properties of the CO2-fixing enzyme Rubisco is a target for improving agricultural crop productivity. Here, we reveal extensive diversity in the kinetic response between 10 and 37 °C by Rubisco from C3 and C4 species within the grass tribe Paniceae. The CO2 fixation rate (kcatc) for Rubisco from the C4 grasses with nicotinamide adenine dinucleotide (NAD) phosphate malic enzyme (NADP-ME) and phosphoenolpyruvate carboxykinase (PCK) photosynthetic pathways was twofold greater than the kcatc of Rubisco from NAD-ME species across all temperatures. The declining response of CO2/O2 specificity with increasing temperature was less pronounced for PCK and NADP-ME Rubisco, which would be advantageous in warmer climates relative to the NAD-ME grasses. Modelled variation in the temperature kinetics of Paniceae C3 Rubisco and PCK Rubisco differentially stimulated C3 photosynthesis relative to tobacco above and below 25 °C under current and elevated CO2. Amino acid substitutions in the large subunit that could account for the catalytic variation among Paniceae Rubisco are identified; however, incompatibilities with Paniceae Rubisco biogenesis in tobacco hindered their mutagenic testing by chloroplast transformation. Circumventing these bioengineering limitations is critical to tailoring the properties of crop Rubisco to suit future climates

C4 photosynthesis boosts growth by altering physiology, allocation and size.

C4 photosynthesis is a complex set of leaf anatomical and biochemical adaptations that have evolved more than 60 times to boost carbon uptake compared with the ancestral C3 photosynthetic type(1-3). Although C4 photosynthesis has the potential to drive faster growth rates(4,5), experiments directly comparing C3 and C4 plants have not shown consistent effects(1,6,7). This is problematic because differential growth is a crucial element of ecological theory(8,9) explaining C4 savannah responses to global change(10,11), and research to increase C3 crop productivity by introducing C4 photosynthesis(12). Here, we resolve this long-standing issue by comparing growth across 382 grass species, accounting for ecological diversity and evolutionary history. C4 photosynthesis causes a 19-88% daily growth enhancement. Unexpectedly, during the critical seedling establishment stage, this enhancement is driven largely by a high ratio of leaf area to mass, rather than fast growth per unit leaf area. C4 leaves have less dense tissues, allowing more leaves to be produced for the same carbon cost. Consequently, C4 plants invest more in roots than C3 species. Our data demonstrate a general suite of functional trait divergences between C3 and C4 species, which simultaneously drive faster growth and greater investment in water and nutrient acquisition, with important ecological and agronomic implications

Contrasting effects of long term versus short-term nitrogen addition on photosynthesis and respiration in the Arctic

We examined the effects of short (<1–4 years) and long-term (22 years) nitrogen (N) and/or phosphorus (P) addition on the foliar CO2 exchange parameters of the Arctic species Betula nana and Eriophorum vaginatum in northern Alaska. Measured variables included: the carboxylation efficiency of Rubisco (Vcmax), electron transport capacity (Jmax), dark respiration (Rd), chlorophyll a and b content (Chl), and total foliar N (N). For both B. nana and E. vaginatum, foliar N increased by 20–50 % as a consequence of 1–22 years of fertilisation, respectively, and for B. nana foliar N increase was consistent throughout the whole canopy. However, despite this large increase in foliar N, no significant changes in Vcmax and Jmax were observed. In contrast, Rd was significantly higher (>25 %) in both species after 22 years of N addition, but not in the shorter-term treatments. Surprisingly, Chl only increased in both species the first year of fertilisation (i.e. the first season of nutrients applied), but not in the longer-term treatments. These results imply that: (1) under current (low) N availability, these Arctic species either already optimize their photosynthetic capacity per leaf area, or are limited by other nutrients; (2) observed increases in Arctic NEE and GPP with increased nutrient availability are caused by structural changes like increased leaf area index, rather than increased foliar photosynthetic capacity and (3) short-term effects (1–4 years) of nutrient addition cannot always be extrapolated to a larger time scale, which emphasizes the importance of long-term ecological experiments