Climatic Drivers of Silicon Accumulation in a Model Grass Operate in Low- but Not High-Silicon Soils.

Grasses are hyper-accumulators of silicon (Si), which is known to alleviate diverse environmental stresses, prompting speculation that Si accumulation evolved in response to unfavourable climatic conditions, including seasonally arid environments. We conducted a common garden experiment using 57 accessions of the model grass Brachypodium distachyon, sourced from different Mediterranean locations, to test relationships between Si accumulation and 19 bioclimatic variables. Plants were grown in soil with either low or high (Si supplemented) levels of bioavailable Si. Si accumulation was negatively correlated with temperature variables (annual mean diurnal temperature range, temperature seasonality, annual temperature range) and precipitation seasonality. Si accumulation was positively correlated with precipitation variables (annual precipitation, precipitation of the driest month and quarter, and precipitation of the warmest quarter). These relationships, however, were only observed in low-Si soils and not in Si-supplemented soils. Our hypothesis that accessions of B. distachyon from seasonally arid conditions have higher Si accumulation was not supported. On the contrary, higher temperatures and lower precipitation regimes were associated with lower Si accumulation. These relationships were decoupled in high-Si soils. These exploratory results suggest that geographical origin and prevailing climatic conditions may play a role in predicting patterns of Si accumulation in grasses.

Field application of silicon alleviates drought stress and improves water use efficiency in wheat.

Detrimental impacts of drought on crop yield have tripled in the last 50 years with climate models predicting that the frequency of such droughts will intensify in the future. Silicon (Si) accumulation, especially in Poaceae crops such as wheat (Triticum aestivum L.), may alleviate the adverse impacts of drought. We have very limited information, however, about whether Si supplementation could alleviate the impacts of drought under field conditions and no studies have specifically manipulated rainfall. Using field-based rain exclusion shelters, we determined whether Si supplementation (equivalent to 39, 78 and 117 kg ha-1) affected T. aestivum growth, elemental chemistry [Si, carbon (C) and nitrogen (N)], physiology (rates of photosynthesis, transpiration, stomatal conductance, and water use efficiency) and yield (grain production) under ambient and drought (50% of ambient) rainfall scenarios. Averaged across Si treatments, drought reduced shoot mass by 21% and grain production by 18%. Si supplementation increased shoot mass by up to 43% and 73% in ambient and drought water treatments, respectively, and restored grain production in droughted plants to levels comparable with plants supplied with ambient rainfall. Si supplementation increased leaf-level water use efficiency by 32-74%, depending on Si supplementation rates. Water supply and Si supplementation did not alter concentrations of C and N, but Si supplementation increased shoot C content by 39% and 83% under ambient and drought conditions, respectively. This equates to an increase from 6.4 to 8.9 tonnes C ha-1 and from 4.03 to 7.35 tonnes C ha-1 under ambient and drought conditions, respectively. We conclude that Si supplementation ameliorated the negative impacts of drought on T. aestivum growth and grain yield, potentially through its beneficial impacts on water use efficiency. Moreover, the beneficial impacts of Si on plant growth and C storage may render Si supplementation a useful tool for both drought mitigation and C sequestration.

Warming drives sustained plant phosphorus demand in a humid tropical forest.

Phosphorus (P) is often one of the most limiting nutrients in highly weathered soils of humid tropical forests and may regulate the responses of carbon (C) feedback to climate warming. However, the response of P to warming at the ecosystem level in tropical forests is not well understood because previous studies have not comprehensively assessed changes in multiple P processes associated with warming. Here, we detected changes in the ecosystem P cycle in response to a 7-year continuous warming experiment by translocating model plant-soil ecosystems across a 600-m elevation gradient, equivalent to a temperature change of 2.1°C. We found that warming increased plant P content (55.4%) and decreased foliar N:P. Increased plant P content was supplied by multiple processes, including enhanced plant P resorption (9.7%), soil P mineralization (15.5% decrease in moderately available organic P), and dissolution (6.8% decrease in iron-bound inorganic P), without changing litter P mineralization and leachate P. These findings suggest that warming sustained plant P demand by increasing the biological and geochemical controls of the plant-soil P-cycle, which has important implications for C fixation in P-deficient and highly productive tropical forests in future warmer climates.

Low phosphorus supply constrains plant responses to elevated CO2 : A meta-analysis.

Phosphorus (P) is an essential macro-nutrient required for plant metabolism and growth. Low P availability could potentially limit plant responses to elevated carbon dioxide (eCO2 ), but consensus has yet to be reached on the extent of this limitation. Here, based on data from experiments that manipulated both CO2 and P for young individuals of woody and non-woody species, we present a meta-analysis of P limitation impacts on plant growth, physiological, and morphological response to eCO2 . We show that low P availability attenuated plant photosynthetic response to eCO2 by approximately one-quarter, leading to a reduced, but still positive photosynthetic response to eCO2 compared to those under high P availability. Furthermore, low P limited plant aboveground, belowground, and total biomass responses to eCO2 , by 14.7%, 14.3%, and 12.4%, respectively, equivalent to an approximate halving of the eCO2 responses observed under high P availability. In comparison, low P availability did not significantly alter the eCO2 -induced changes in plant tissue nutrient concentration, suggesting tissue nutrient flexibility is an important mechanism allowing biomass response to eCO2 under low P availability. Low P significantly reduced the eCO2 -induced increase in leaf area by 14.3%, mirroring the aboveground biomass response, but low P did not affect the eCO2 -induced increase in root length. Woody plants exhibited stronger attenuation effect of low P on aboveground biomass response to eCO2 than non-woody plants, while plants with different mycorrhizal associations showed similar responses to low P and eCO2 interaction. This meta-analysis highlights crucial data gaps in capturing plant responses to eCO2 and low P availability. Field-based experiments with longer-term exposure of both CO2 and P manipulations are critically needed to provide ecosystem-scale understanding. Taken together, our results provide a quantitative baseline to constrain model-based hypotheses of plant responses to eCO2 under P limitation, thereby improving projections of future global change impacts.

Effects of elevated carbon dioxide and elevated temperature on morphological, physiological and anatomical responses of Eucalyptus tereticornis along a soil phosphorus gradient.

Eucalypts are likely to play a critical role in the response of Australian forests to rising atmospheric CO2 concentration ([CO2]) and temperature. Although eucalypts are frequently phosphorus (P) limited in native soils, few studies have examined the main and interactive effects of P availability, [CO2] and temperature on eucalypt morphology, physiology and anatomy. To address this issue, we grew seedlings of Eucalyptus tereticornis Smith across its P-responsive range (6-500 mg kg-1) for 120 days under two [CO2] (ambient: 400 µmol mol-1 (Ca) and elevated: 640 µmol mol-1 (Ce)) and two temperature (ambient: 24/16 °C (Ta) and elevated: 28/20 °C (Te) day/night) treatments in a sunlit glasshouse. Seedlings were well-watered and supplied with otherwise non-limiting macro- and micro-nutrients. Increasing soil P supply increased growth responses to Ce and Te. At the highest P supplies, Ce increased total dry mass, leaf number and total leaf area by ~50%, and Te increased leaf number by ~40%. By contrast, Ce and Te had limited effects on seedling growth at the lowest P supply. Soil P supply did not consistently modify photosynthetic responses to Ce or Te. Overall, effects of Ce and Te on growth, physiological and anatomical responses of E. tereticornis seedlings were generally neutral or negative at low soil P supply, suggesting that native tree responses to future climates may be relatively small in native low-P soils in Australian forests.

Soil phosphorous and endogenous rhythms exert a larger impact than CO2 or temperature on nocturnal stomatal conductance in Eucalyptus tereticornis.

High nocturnal transpiration rates (5-15% of total water loss in terrestrial plants) may be adaptive under limited fertility, by increasing nutrient uptake or transport via transpiration-induced mass flow, but the response of stomata in the dark to environmental variables is poorly understood. Here we tested the impact of soil phosphorous (P) concentration, atmospheric CO2 concentration and air temperature on stomatal conductance (gs) during early and late periods in the night, as well as at midday in naturally, sun-lit glasshouse-grown Eucalyptus tereticornis Sm. seedlings. Soil P was the main driver of nocturnal gs, which was consistently higher in low soil P (37.3-79.9 mmol m(-2) s(-1)) than in high soil P (17.7-49.3 mmol m(-2)(-1)). Elevated temperature had only a marginal (P = 0.07) effect on gs early in the night (gs decreased from 34.7 to 25.8 mmol m(-2) s(-1) with an increase in temperature of 4 °C). The effect of CO2 depended on its interaction with temperature. Stomatal conductance responses to soil P were apparently driven by indirect effects of soil P on plant anatomy, since gs was significantly and negatively correlated with wood density. However, the relationship of gs with environmental factors became weaker late in the night, relative to early in the night, likely due to apparent endogenous processes; gs late in the night was two times larger than gs observed early in the night. Time-dependent controls over nocturnal gs suggest that daytime stomatal models may not apply during the night, and that different types of regulation may occur even within a single night. We conclude that the enhancement of nocturnal gs under low soil P availability is unlikely to be adaptive in our species because of the relatively small amount of transpiration-induced mass flow that can be achieved through rates of nocturnal water loss (3-6% of daytime mass flow).

Impacts of drought on leaf respiration in darkness and light in Eucalyptus saligna exposed to industrial-age atmospheric CO2 and growth temperature.

Our study assessed the impact of a wide range of industrial-age climate scenarios on leaf respiration (R) in Eucalyptus saligna. Well-watered or sustained drought-treated plants were grown in glasshouses differing in atmospheric CO2 concentration ([CO2]) (280, 400 and 640 µl l⁻¹) and temperature (26 and 30°C). Rates of R in darkness (R(dark) ) and light (R(light) ), photosynthesis (A) and related leaf traits (mass : area relationships, and nitrogen, phosphorus, starch and sugar concentrations) were measured. Light inhibited R in all cases (R(light) < R(dark) ) (well-watered: 40%; drought-treated: 73%). Growth [CO2] and temperature had little impact on area-based rates of R(dark) or R(light) , with R(light) exhibiting minimal thermal acclimation. By contrast, sustained drought resulted in reduced R(dark), R(light) and A, with the inhibitory effect of drought on A and R(light) (c. 50-70%) greater than that on R(dark) (c. 15%). Drought effects were fully reversible after watering. Variability in R(light) appeared to be dependent on the underlying rate of R(dark) and associated Rubisco activity. Collectively, our data suggest that there is an asynchronous response of leaf carbon metabolism to drought, and a tighter coupling between R(light) and A than between R(dark) and A, under both past and future climate scenarios. These findings have important implications for ecosystem/global models seeking to predict carbon cycling.

Photosynthetic responses of cottonwood seedlings grown in glacial through future atmospheric [CO2] vary with phosphorus supply.

Plants often exhibit proportionately larger photosynthetic responses to the transition from glacial to modern [CO(2)] than from modern to future [CO(2)]. Although this pattern may reflect increased nutrient demand with increasing [CO(2)], few studies have examined the role of nutrient supply in regulating responses to the range of [CO(2)] from glacial to future [CO(2)]. In this study, we examined the effects of P supply (0.004-0.5 mM) on photosynthetic responses of Populus deltoides (cottonwood) seedlings to glacial (200 micromol mol(-1)), modern (350 µmol mol(-1)) and future (700 micromol mol(-1)) [CO(2)]. The A(sat) (light-saturated net photosynthetic rates at the growth [CO(2)]) response to future [CO(2)] decreased with decreasing P supply such that there was no response at the lowest P supply. However, P supply did not affect A(sat) responses to an increase from glacial to modern [CO(2)]. Photosynthetic capacity [e.g., final rubisco activity, apparent, maximal Rubisco-limited rate of photosynthesis (V(cmax)), apparent, maximal electron transport-limited rate of photosynthesis (J(max))], stomatal conductance (g(s)) and leaf P generally increased with increasing P supply but decreased with increasing [CO(2)]. Measures of carbohydrate sink capacity (e.g., leaf mass per unit leaf area, leaf starch) increased with both increasing P supply and increasing [CO(2)]. Changes in V(cmax) and g(s) together accounted for 78% of the variation in A(sat) among [CO(2)] and P treatments, suggesting significant biochemical and stomatal controls on photosynthesis. However, A(sat) responses to increasing [CO(2)] did not reflect the changes in the carbohydrate sink capacity. These results have important implications because low P already constrains responses to increasing [CO(2)] in many ecosystems, and our results suggest that the P demand will increasingly affect A(sat) in cottonwood as [CO(2)] continues to increase.

Phosphorus supply drives nonlinear responses of cottonwood (Populus deltoides) to increases in CO2 concentration from glacial to future concentrations.

SUMMARY: *Despite the importance of nutrient availability in determining plant responses to climate change, few studies have addressed the interactive effects of phosphorus (P) supply and rising atmospheric CO(2) concentration ([CO(2)]) from glacial to modern and future concentrations on tree seedling growth. *The objective of our study was to examine interactive effects across a range of P supply (six concentrations from 0.004 to 0.5 mM) and [CO(2)] (200 (glacial), 350 (modern) and 700 (future) ppm) on growth, dry mass allocation, and light-saturated photosynthesis (A(sat)) in Populus deltoides (cottonwood) seedlings grown in well-watered conditions. *Increasing [CO(2)] from glacial to modern concentrations increased growth by 25% across P treatments, reflecting reduced [CO(2)] limitations to photosynthesis and increased A(sat). Conversely, the growth response to future [CO(2)] was very sensitive to P supply. Future [CO(2)] increased growth by 80% in the highest P supply but only by 7% in the lowest P supply, reflecting P limitations to A(sat), leaf area and leaf area ratio (LAR), compared with modern [CO(2)]. *Our results suggest that future [CO(2)] will minimally increase cottonwood growth in low-P soils, but in high-P soils may stimulate production to a greater extent than predicted based on responses to past increases in [CO(2)]. Our results indicate that the capacity for [CO(2)] stimulation of cottonwood growth does not decline as [CO(2)] rises from glacial to future concentrations.

A hierarchical Bayesian approach for estimation of photosynthetic parameters of C(3) plants.

We describe a hierarchical Bayesian (HB) approach to fitting the Farquhar et al.model of photosynthesis to leaf gas exchange data. We illustrate the utility of this approach for estimating photosynthetic parameters using data from desert shrubs. Unique to the HB method is its ability to simultaneously estimate plant- and species-level parameters, adjust for peaked or non-peaked temperature dependence of parameters, explicitly estimate the 'critical' intracellular [CO(2)] marking the transition between ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco) and ribulose-1,5-bisphosphate (RuBP) limitations, and use both light response and CO(2) response curve data to better inform parameter estimates. The model successfully predicted observed photosynthesis and yielded estimates of photosynthetic parameters and their uncertainty. The model with peaked temperature responses fit the data best, and inclusion of light response data improved estimates for day respiration (R(d)). Species differed in R(d25) (R(d) at 25 degrees C), maximum rate of electron transport (J(max25)), a Michaelis-Menten constant (K(c25)) and a temperature dependence parameter (DeltaS). Such differences could potentially reflect differential physiological adaptations to environmental variation. Plants differed in R(d25), J(max25), mesophyll conductance (g(m25)) and maximum rate of Rubisco carboxylation (V(cmax25)). These results suggest that plant- and species-level variation should be accounted for when applying the Farquhar et al. model in an inferential or predictive framework.

Photosynthesis and reflectance indices for rainforest species in ecosystems undergoing progression and retrogression along a soil fertility chronosequence in New Zealand.

Measurements of photosynthesis at saturating irradiance and CO2 partial pressure, Amax, "adjusted" normalised difference vegetation index, RaNDVI, and photochemical reflectance index, RPRI, were made on trees sampled along a soil chronosequence to investigate the relationship between carbon uptake and ecosystem development in relation to nutrient availability. Measurements were made on the three most dominant species at six sites along the sequence in South Westland, New Zealand with soil age ranging from < 6 to 120,000 years resulting from the retreat of the Franz Josef glacier. The decrease in soil phosphorus availability with increasing soil age and high soil nitrogen availability at the two youngest sites, due to the presence of a nitrogen-fixing species, provided marked differences in nutrient availability. Mean Amax was high at the two youngest sites, then decreased markedly with increasing site age. Analysis of the data for individual species within sites revealed separation of groups of species in the response of Amax to Nm and Pm, suggesting complex interactions between the two nutrients. There were strong linear relationships for leaf-level RaNDVI and RPRI with Amax, at high irradiance, showing that measurements of reflectance indices can be used to estimate Amax for foliage with a range in morphology and nutrient concentrations. Notwithstanding the change in species composition from angiosperms to conifers with increasing site age, the presence of nitrogen-fixing species, the variability in foliage morphology from flat leaves to imbricate scales and a wide range in foliar nitrogen and phosphorus concentrations, there were strong positive linear relationships between site average Amax and foliage nitrogen, Nm, and phosphorus, Pm, concentrations on a foliage mass basis. The results provide insights to interpret the regulation of photosynthesis across natural ecosystems with marked gradients in nitrogen and phosphorus availability.

Atmospheric CO2 enrichment alters energy assimilation, investment and allocation in Xanthium strumarium.

Energy-use efficiency and energy assimilation, investment and allocation patterns are likely to influence plant growth responses to increasing atmospheric CO2 concentration ([CO2]). Here, we describe the influence of elevated [CO2] on energetic properties as a mechanism of growth responses in Xanthium strumarium. Individuals of X. strumarium were grown at ambient or elevated [CO2] and harvested. Total biomass and energetic construction costs (CC) of leaves, stems, roots and fruits and percentage of total biomass and energy allocated to these components were determined. Photosynthetic energy-use efficiency (PEUE) was calculated as the ratio of total energy gained via photosynthetic activity (Atotal) to leaf CC. Elevated [CO2] increased leaf Atotal, but decreased CC per unit mass of leaves and roots. Consequently, X. strumarium individuals produced more leaf and root biomass at elevated [CO2] without increasing total energy investment in these structures (CCtotal). Whole-plant biomass was associated positively with PEUE. Whole-plant construction required 16.1% less energy than modeled whole-plant energy investment had CC not responded to increased [CO2]. As a physiological mechanism affecting growth, altered energetic properties could positively influence productivity of X. strumarium, and potentially other species, at elevated [CO2].

Stomatal and non-stomatal limitations to photosynthesis in four tree species in a temperate rainforest dominated by Dacrydium cupressinum in New Zealand.

We assessed the relative limitations to photosynthesis imposed by stomatal and non-stomatal processes in Dacrydium cupressinum Lamb. (Podocarpaceae), which is the dominant species in a native, mixed conifer-broad-leaved rainforest in New Zealand. For comparison, we included three co-occurring broad-leaved tree species (Meterosideros umbellata Cav. (Myrtaceae), Weinmannia racemosa L.f. (Cunoniaceae) and Quintinia acutifolia Kirk (Escalloniaceae)) that differ in phylogeny and in leaf morphology from D. cupressinum. We found that low foliage phosphorus content on an area basis (P(a)) limited light-saturated photosynthesis on an area basis (A(sat)) in Q. acutifolia. Depth in the canopy did not generally affect A(sat) or the relative limitations to A(sat) because of stomatal and non-stomatal constraints, despite reductions in the ratio of foliage mass to area, foliar nitrogen on an area basis (N(a)) and P(a) with depth in the canopy. In the canopy-dominant conifer D. cupressinum, A(sat) was low, consistent with low values of the maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V(cmax)). In comparison, the A(sat) response of the three broad-leaved tree species was quite variable. Although A(sat) was high in the canopy-dominant M. umbellata, it was low in the sub-canopy trees W. racemosa and Q. acutifolia. Relative stomatal limitation to photosynthesis was more pronounced in W. racemosa (40%) than in the other three species (28-33%). Despite differences in degree, non-stomatal limitation to A(sat) predominated in all tree species.

Respiration characteristics in temperate rainforest tree species differ along a long-term soil-development chronosequence.

We measured the response of dark respiration (R(d)) to temperature and foliage characteristics in the upper canopies of tree species in temperate rainforest communities in New Zealand along a soil chronosequence (six sites from 6 years to 120,000 years). The chronosequence provided a vegetation gradient characterised by significant changes in soil nutrition. This enabled us to examine the extent to which changes in dark respiration can be applied across forest biomes and the utility of scaling rules in whole-canopy carbon modelling. The response of respiration to temperature in the dominant tree species differed significantly between sites along the sequence. This involved changes in both R(d) at a reference temperature (R(10)) and the extent to which R(d) increased with temperature (described by E(o), a parameter related to the energy of activation, or the change in R(d) over a 10 degrees C range, Q(10)). Site averaged E(o) ranged from 44.4 kJ mol(-1) K(-1) at the 60-year-old site to 26.0 kJ mol(-1) K(-1) at the oldest, most nutrient poor, site. Relationships between respiratory and foliage characteristics indicated that both the temperature response of respiration (E(o) or Q(10)) and the instantaneous rate of respiration increased with both foliar nitrogen and phosphorus content. The ratio of photosynthetic capacity (Whitehead et al. in Oecologia 2005) to respiration (A(max)/R(d)) attained values in excess of 15 for species in the 6- to 120-year-old sites, but thereafter decreased significantly to around five at the 120,000-year-old site. This indicates that shoot carbon acquisition is regulated by nutrient limitations in the retrogressing ecosystems on the oldest sites. Our findings indicate that respiration and its temperature response will vary according to soil age and, therefore, to soil nutrient availability and the stage of forest development. Thus, variability in respiratory characteristics for canopies should be considered when using models to integrate respiration at large spatial scales.

Age at flowering differentially affects vegetative and reproductive responses of a determinate annual plant to elevated carbon dioxide.

Plant population and community dynamics may be altered by increasing atmospheric CO(2) concentrations [[CO(2)]] through intraspecific variation in the responses of vegetative and reproductive growth. Although these responses may be regulated by age at flowering, little is known about the direct effects of age at flowering on growth responses to elevated [CO(2)]. In this study, we examined the interactive effects of elevated [CO(2)] and age at flowering on absolute and relative allocation to vegetative and reproductive growth in the determinate, short-day species Xanthium strumarium L. (common cocklebur). Six cohorts were planted at 5-day intervals in chambers maintained at either 365 or 730 micro mol mol(-1) CO(2), with an 18-h photoperiod and a non-limiting nutrient supply. All plants were simultaneously induced to flower by switching the photoperiod to 12 h for 2 days, then switching back to an 18-h photoperiod for the remainder of the experiment. All plants were harvested 15 days after the onset of flowering. Total plant biomass increased 11-41% with increasing [CO(2)] and 45% from the youngest to the oldest cohort. Vegetative growth responses to elevated [CO(2)] significantly increased with increasing age at flowering, associated with increasing sink relative to source capacity. In contrast, total fruit mass decreased 32% from the youngest to the oldest cohort and was not significantly affected by CO(2) supply. Relative biomass allocation to fruit decreased 47% from the youngest to the oldest cohort, reflecting decreased numbers of fruit, and 6-28% with increasing [CO(2)], reflecting decreased mean mass per mature fruit. Our findings suggest that elevated [CO(2)] may increase vegetative growth in Xanthium without increasing reproductive biomass, and that age at flowering may influence these responses through effects on source:sink balance. Further, changes in the allometric relationship between vegetative and reproductive growth associated with growth in elevated [CO(2)] suggest that long-term population and community-level responses to elevated [CO(2)] may differ substantially from predictions based on vegetative responses.