Matthew H. Turnbull

The effect of light quantity and quality during development on the photosynthetic characteristics of six Australian rainforest tree species

SummarySeedlings of six subtropical rainforest tree species representing early (Omalanthus populifolius, Solanum aviculare), middle (Duboisia myoporoides, Euodia micrococca) and late (Acmena ingens, Argyrodendron actinophyllum) successional stages in forest development were grown in a glasshouse, under four levels of neutral shade (60%, 15%, 5%, 1% of photosynthetically active radiation (PAR) in incident sunlight) and three levels of selectively filtered shade (producing 15%, 5%, 1% of PAR). This design served to analyse the interactions between reduced photon flux density (PFD) and reduced red/far-red (R/FR) ratio in their effects on selected photosynthetic characteristics of each species. The light-saturated rate of photosynthesis was significantly influenced by growth irradiance in five of the six species, with all of these showing a non-linear decrease in maximum assimilation rate from 60% down to 1% PAR. The degree of acclimation to this range was not clearly related to the successional status of the species. Dark respiration was more sensitive to growth irradiance in the early- and mid-stage species than in the late-stage species. Although levels of dark respiration were clearly greater in leaves of early- and mid-stage species from the highest light levels, differences between successional groups were negligible at 1% PAR. Growth in filtered shade, typical of that beneath a closed canopy, resulted in lower photosynthetic capacities and quantum yields in those species which did respond. Although dark respiration rates were more sensitive to filtered shade in the early-stage than in the late-stage species, there was no evidence from other gas exchange characteristics to suggest that overall sensitivity to light quality (as characterised by the R/FR ratio) is greater in early successional-stage species.

Plant growth in elevated CO2 alters mitochondrial number and chloroplast fine structure.

With increasing interest in the effects of elevated atmospheric CO2 on plant growth and the global carbon balance, there is a need for greater understanding of how plants respond to variations in atmospheric partial pressure of CO2. Our research shows that elevated CO2 produces significant fine structural changes in major cellular organelles that appear to be an important component of the metabolic responses of plants to this global change. Nine species (representing seven plant families) in several experimental facilities with different CO2-dosing technologies were examined. Growth in elevated CO2 increased numbers of mitochondria per unit cell area by 1.3–2.4 times the number in control plants grown in lower CO2 and produced a statistically significant increase in the amount of chloroplast stroma (nonappressed) thylakoid membranes compared with those in lower CO2 treatments. There was no observable change in size of the mitochondria. However, in contrast to the CO2 effect on mitochondrial number, elevated CO2 promoted a decrease in the rate of mass-based dark respiration. These changes may reflect a major shift in plant metabolism and energy balance that may help to explain enhanced plant productivity in response to elevated atmospheric CO2 concentrations.

The dynamics of photosynthetic acclimation to changes in light quanlity and quality in three Australian rainforest tree species

Photosynthetic acclimation was studied in seedlings of three subtropical rainforest species representing early (Omalanthus populifolius), middle (Duboisia myoporoides) and late (Acmena ingens) successional stages in forest development. Changes in the photosynthetic characteristics of pre-existing leaves were observed following the transfer of plants between deep shade (1–5% of photosynthetically active radiation (PAR), selectively filtered to produce a red/far-red (R/FR) ratio of 0.1) and open glasshouse (60% PAR and a R/FR ratio of 1.1–1.2), and vice versa. The extent and rate of response of the photosynthetic characteristics of each species to changes in light environment were recorded in this simulation of gap formation and canopy closure/overtopping. The light regimes to which plants were exposed produced significant levels of acclimation in all the photosynthetic parameters examined. Following transfer from high to low light, the light-saturated rate of photosynthesis was maintained near pre-transfer levels for 7 days, after which it decreased to levels which closely approximated those in leaves which had developed in low light. The decrease in photosynthetic capacity was associated with lower apparent quantum yields and stomatal conductances. Dark respiration was the parameter most sensitive to changes in light environment, and responded significantly during the first 4–7 days after transfer. Acclimation of photosynthetic capacity to increases in irradiance was significant in two of the three species studied, but was clearly limited in comparison with that of new leaves produced in the high light conditions. This limitation was most pronounced in the early-successional-stage species, O. populifolius. It is likely that structural characteristics of the leaves, imposed at the time of leaf expansion, are largely responsible for the limitations in photosynthetic acclimation to increases in irradiance.

Evidence That Glutamate Dehydrogenase Plays a Role in the Oxidative Deamination of Glutamate in Seedlings of Zea mays

In order to investigate the role of glutamate dehydrogenase we have compared the metabolism of [15N]glutamate in young seedlings of wild-type and a glutamate dehydrogenase-null mutant of Zea mays. The principal labelled products in roots of wild-type seedlings are the amide nitrogen of glutamine, glutamine-amino nitrogen and ammonium. The incorporation of label into glutamine-amide is markedly inhibited by methionine sulfoximine. In contrast, little or no labelling of glutamine-amide or ammonium occurs in roots of the GDH1-null mutant, the major labelled product is the amino group of asparagine. In shoots of the wild type, 15N is recovered in the amide of glutamine, ammonium, the amino group of asparagine and other amino acids. In mutant shoots, over 75% of the label is recovered in the asparagine-amino group and there is little labelling of glutamine-amide or ammonium. These major differences in glutamate metabolism of wild-type and mutant seedlings are consistent with glutamate dehydrogenase functioning in the direction of oxidative deamination and having a role in protein catabolism of germinating seeds.

Increased pond depth improves algal productivity and nutrient removal in wastewater treatment high rate algal ponds

Depth has been widely recognised as a crucial operational feature of a high rate algal pond (HRAP) as it modifies the amount of light and frequency at which microalgal cells are exposed to optimal light. To date, there has been little focus on the optimisation of microalgal performance in wastewater treatment HRAPs with respect to depth, with advice ranging from as shallow as possible to 100 cm deep. This paper investigates the seasonal performance of microalgae in wastewater treatment HRAPs operated at three different depths (200, 300 and 400 mm). Microalgal performance was measured in terms of biomass production and areal productivity, nutrient removal efficiency and photosynthetic performance. The overall areal productivity significantly increased with increasing depth. Areal productivity ranged from 134 to 200% higher in the 400 mm deep HRAP compared to the 200 mm deep HRAP. Microalgae in the 400 mm deep HRAP were more efficient at NH4-N uptake and were photosynthetically more efficient compared to microalgae in the 200 mm deep HRAP. A higher chlorophyll-a concentration in the 200 mm deep HRAP resulted in a decrease in photosynthetic performance, due to insufficient carbon supply, over the course of the day in summer (as indicated by lower α, Pmax and oxygen production) compared to the 300 and 400 mm deep HRAPs. Based on these results, improved areal productivity and more wastewater can be treated per land area in the 400 mm deep HRAPs compared to 200 mm deep HRAPs without compromising wastewater treatment quality, while lowering capital and operational costs.

Nocturnal stomatal conductance and implications for modelling δ18O of leaf-respired CO2 in temperate tree species

Variation in the oxygen isotope composition of within-canopy CO2 has potential to allow partitioning of the ecosystem respiratory flux into above- and below-ground components. Recent theoretical work has highlighted the sensitivity of the oxygen isotope composition of leaf-respired CO2 (δRl) to nocturnal stomatal conductance. When the one-way flux model was tested on Ricinus communis L. large enrichments in δRl were observed. However, most species for which the isotope flux partitioning technique has been or would be applied (i.e. temperate tree species) are much more conservative users of water than R. communis. So, high stomatal conductance and very high enrichment of δRl observed may not be typical for temperate tree species. Using existing gas-exchange measurements on six temperate tree species, we demonstrate significant water loss through stomata for all species (i.e. statistically significantly greater than cuticular loss alone) at some time for some leaves during the night. δRl values predicted by the one-way flux model revealed that δRl might be very much more enriched than when the net flux alone is considered, particularly close to sunrise and sunset. Incorporation of the one-way flux model into ecosystem respiration partitioning studies will affect model outputs and interpretation of variation in the oxygen isotope composition of atmospheric CO2.

Bringing the Kok effect to light: A review on the integration of daytime respiration and net ecosystem exchange

Net ecosystem exchange (NEE) represents the difference between carbon assimilated through photosynthesis, or gross primary productivity (GPP), and carbon released via ecosystem respiration (ER). NEE, measured via eddy covariance and chamber techniques, must be partitioned into these fluxes to accurately describe and understand the carbon dynamics of an ecosystem. GPP and daytime ER may be significantly overestimated if the light inhibition of foliar mitochondrial respiration, or “Kok effect,” is not accurately estimated and further integrated into ecosystem measurements. The light inhibition of respiration, a composite effect of multiple cellular pathways, is reported to cause between 25-100% inhibition of foliar mitochondrial respiration, and for this reason needs to be considered when estimating larger carbon fluxes. Partitioning of respiration between autotrophic and heterotrophic respiration, and applying these scaled respiratory fluxes to the ecosystem-level proves to be difficult, and the integration of light inhibition into single and continuous measures of ecosystem respiration will require new interpretations and analysis of carbon exchange in terrestrial ecosystems.

Leaf respiration and alternative oxidase in field‐grown alpine grasses respond to natural changes in temperature and light

• We report the first investigation of changes in electron partitioning via the alternative respiratory pathway (AP) and alternative oxidase (AOX) protein abundance in field-grown plants and their role in seasonal acclimation of respiration. • We sampled two alpine grasses native to New Zealand, Chionochloa rubra and Chionochloa pallens, from field sites of different altitudes, over 1 yr and also intensively over a 2-wk period. • In both species, respiration acclimated to seasonal changes in temperature through changes in basal capacity (R₁₀) but not temperature sensitivity (E₀). In C. pallens, acclimation of respiration may be associated with a higher AOX : cytochrome c oxidase (COX) protein abundance ratio. Oxygen isotope discrimination (D), which reflects relative changes in AP electron partitioning, correlated positively with daily integrated photosynthetically active radiation (PAR) in both species over seasonal timescales. Respiratory parameters, the AOX : COX protein ratio and D were stable over a 2-wk period, during which significant temperature changes were experienced in the field. • We conclude that respiration in Chionochloa spp. acclimates strongly to seasonal, but not to short-term, temperature variation. Alternative oxidase appears to be involved in the plant response to both seasonal changes in temperature and daily changes in light, highlighting the complexity of the function of AOX in the field.

Convergence in the temperature response of leaf respiration across biomes and plant functional types.

Significance A major concern for terrestrial biosphere models is accounting for the temperature response of leaf respiration at regional/global scales. Most biosphere models incorrectly assume that respiration increases exponentially with rising temperature, with profound effects for predicted ecosystem carbon exchange. Based on a study of 231 species in 7 biomes, we find that the rise in respiration with temperature can be generalized across biomes and plant types, with temperature sensitivity declining as leaves warm. This finding indicates universally conserved controls on the temperature sensitivity of leaf metabolism. Accounting for the temperature function markedly lowers simulated respiration rates in cold biomes, which has important consequences for estimates of carbon storage in vegetation, predicted concentrations of atmospheric carbon dioxide, and future surface temperatures. Plant respiration constitutes a massive carbon flux to the atmosphere, and a major control on the evolution of the global carbon cycle. It therefore has the potential to modulate levels of climate change due to the human burning of fossil fuels. Neither current physiological nor terrestrial biosphere models adequately describe its short-term temperature response, and even minor differences in the shape of the response curve can significantly impact estimates of ecosystem carbon release and/or storage. Given this, it is critical to establish whether there are predictable patterns in the shape of the respiration–temperature response curve, and thus in the intrinsic temperature sensitivity of respiration across the globe. Analyzing measurements in a comprehensive database for 231 species spanning 7 biomes, we demonstrate that temperature-dependent increases in leaf respiration do not follow a commonly used exponential function. Instead, we find a decelerating function as leaves warm, reflecting a declining sensitivity to higher temperatures that is remarkably uniform across all biomes and plant functional types. Such convergence in the temperature sensitivity of leaf respiration suggests that there are universally applicable controls on the temperature response of plant energy metabolism, such that a single new function can predict the temperature dependence of leaf respiration for global vegetation. This simple function enables straightforward description of plant respiration in the land-surface components of coupled earth system models. Our cross-biome analyses shows significant implications for such fluxes in cold climates, generally projecting lower values compared with previous estimates.

Canopy position affects the relationships between leaf respiration and associated traits in a tropical rainforest in Far North Queensland

We explored the impact of canopy position on leaf respiration (R) and associated traits in tree and shrub species growing in a lowland tropical rainforest in Far North Queensland, Australia. The range of traits quantified included: leaf R in darkness (RD) and in the light (RL; estimated using the Kok method); the temperature (T)-sensitivity of RD; light-saturated photosynthesis (Asat); leaf dry mass per unit area (LMA); and concentrations of leaf nitrogen (N), phosphorus (P), soluble sugars and starch. We found that LMA, and area-based N, P, sugars and starch concentrations were all higher in sun-exposed/upper canopy leaves, compared with their shaded/lower canopy and deep-shade/understory counterparts; similarly, area-based rates of RD, RL and Asat (at 28 °C) were all higher in the upper canopy leaves, indicating higher metabolic capacity in the upper canopy. The extent to which light inhibited R did not differ significantly between upper and lower canopy leaves, with the overall average inhibition being 32% across both canopy levels. Log-log RD-Asat relationships differed between upper and lower canopy leaves, with upper canopy leaves exhibiting higher rates of RD for a given Asat (both on an area and mass basis), as well as higher mass-based rates of RD for a given [N] and [P]. Over the 25-45 °C range, the T-sensitivity of RD was similar in upper and lower canopy leaves, with both canopy positions exhibiting Q10 values near 2.0 (i.e., doubling for every 10 °C rise in T) and Tmax values near 60 °C (i.e., T where RD reached maximal values). Thus, while rates of RD at 28 °C decreased with increasing depth in the canopy, the T-dependence of RD remained constant; these findings have important implications for vegetation-climate models that seek to predict carbon fluxes between tropical lowland rainforests and the atmosphere.