Nitrogen allocation in response to partial shading of a plant: Possible mechanisms

The significance of photosynthetic and transpiration rates for the perception by plants of light gradients in leaf canopies has been investigated with regard to nitrogen allocation and re-allocation. A gradient of photon flux density (PFD) over a plant's foliage was simulated by shading one leaf of a pair of primary leaves of bean ( Phaseolus vulgaris L. cv. Rentegever). Photosynthetic rate was manipulated independently of PFD and, to some extent, also of transpiration, by subjecting the leaf to different CO₂ concentrations. Transpiration rate was changed independently of PFD and photosynthetic rate by subjecting the leaf to different vapour pressure differences (VPD). A reduced partial pressure of CO₂ reduced specific leaf mass (SLM) as did a decreased PFD, but did not change leaf N per unit area (N_LA) and light saturated rate of photosynthesis (A_max). A reduced VPD caused several effects consistent with the effect of PFD. It decreased N_LA and A_max and increased the chlorophyll to N ratio in old and young leaves. Furthermore, it decreased the chlorophyll a to b ratio and inhibited leaf growth in young leaves. The transpiration stream is partitioned among the leaves of a plant according to their transpiration rates. The results suggest that relative rates of import of xylem sap into leaves of a plant play an important role in the perception of partial shading of a plant, a situation normally found in dense vegetations. The possible role of cytokinin influx into leaves as controlled by transpiration rate, is discussed.     

Nitrogen reallocation and photosynthetic acclimation in response to partial shading in soybean plants

The first trifoliate of soybean was shaded when fully expanded, while the plant remained in high light; a situation representative for plants growing in a closed crop. Leaf mass and respiration rate per unit area declined sharply in the first few days upon shading and remained rather constant during the further 12 days of the shading treatment. Leaf nitrogen per unit area decreased gradually until the leaves were shed. Leaf senescence was enhanced by the shading treatment in contrast to control plants growing in low light. Shaded leaves on plants grown at low nutrient availability senesced earlier than shaded leaves on plants grown at high nutrient availability. The light saturated rate of photosynthesis decreased also gradually during the shading treatment, but somewhat faster than leaf N, whereas chlorophyll contents declined somewhat slower than leaf N.
Partitioning of N in the leaf over main photosynthetic functions was estimated from parameters derived from the response of photosynthesis to CO₂. It appeared that the N exported from the leaf was more at the expense of compounds that make up photosynthetic capacity than of those involved in photon absorption, resulting in a change in partitioning of N within the photosynthetic apparatus. Photosynthetic nitrogen use efficiency increased during the shading treatment, which was for the largest part due to the decrease in leaf N content, to some extent to the decrease in respiration rate and only for a small part to change in partitioning of N within the photosynthetic apparatus.     

羊草种群对遮荫的适应性生长对策

自20世纪50年代以来,太阳到达整个地球的短波光能辐射量每10年减少2.7％,即“全球变暗”,这会对植物及其所在的生态系统的功能产生影响,然而,目前关于这种影响的研究尚未得出一致的结论.本文利用不同程度的遮荫(0、30％、50％和70％)模拟全球变暗的效应,研究了我国北方草原优势植物羊草的种群密度、形态指标和光合生理等对遮荫的响应.结果表明:遮荫使羊草枝条密度有所降低,枝条高度明显增加,叶面积显著扩大,抽穗枝条比例即抽穗率升高;遮荫使羊草接受到的光合有效辐射显著下降,在0、30％、50％遮荫强度下羊草的气孔导度、光合速率和蒸腾速率仍表现出“午休”现象,而遮荫70％的处理下气孔导度、光合和蒸腾速率在11:00-16:00时间段均高于其他处理,而且光合和蒸腾速率未出现“午休”现象.这种“午休”现象的消失可能是一定程度遮荫或全球变暗导致某些植被系统生产力增加的重要原因之一.为更加准确预测各类生态系统对全球变暗的响应,必须针对更多地域、更多生境、更多类群和更多生育期植物对遮荫的响应开展实验研究,并加以整合和模拟.     

Effects of environment during growth on the sensitivity of leaf conductance to changes in humidity

Soybeans (Glycine max) and grain amaranth (Amaranthus hypochondriacus) were grown at a range of temperatures, carbon dioxide concentrations and light conditions in controlled environment chambers, and the response of leaf conductance to water vapour to changes in humidity was then measured under a standard set of conditions. The sensitivity of conductance was analysed in terms of (i) the absolute sensitivity of conductance to changes in leaf to air water vapour pressure difference (LAVPD), (ii) the sensitivity of conductance relative to the absolute value of conductance, and (iii) the slope of the relationship between conductance and an index incorporating assimilation rate, carbon dioxide concentration and relative humidity. The sensitivity of conductance varied substantially with growth conditions for all three analyses in both species. The growth temperature of 25 °C increased the sensitivity of conductance by all three measures compared with growth at 20 or 30 °C in amaranth, with little difference between 25 and 30 °C in soybean. Growth at elevated carbon dioxide decreased sensitivity in amaranth by all three measures, and decreased the absolute but not the relative sensitivity in soybean. Growth at reduced photon flux density and growth at high stand density reduced sensitivity in amaranth by all three measures. In soybean, growth at high stand density reduced sensitivity by all three measures, but growth at low photon flux density increased the relative sensitivity. The sensitivity of leaf conductance to changes in humidity varied by a factor of two or more with growth environment by all measures of sensitivity in both the C3 and the C4 species.     

Stomatal response to humidity in a sugarcane field: simultaneous porometric and micrometeorological measurements*

Abstract. Gas exchange data obtained with wellventilated leaf cuvettes provide clear evidence of a stomatal response to leaf-air vapour pressure difference (V). In contrast, remotely sensed leaf temperatures with specific assumptions regarding canopy boundary layer characteristics, have been interpreted to mean that stomata do not respond to V. We address this apparent discrepancy in a sugarcane field by simultaneous application of a single-leaf, porometric technique and a whole-canopy, Bowen ratioenergy balance technique. These methods indicated significant stomatal response to V in well-irrigated sugarcane. Stomatal responses to V in the field were obscured by strong covariance of major environmental parameters so that opening responses to light and closing responses to V tended to offset each other. Low boundary layer conductance significantly uncoupled V at the leaf surface (V_s) from V determined in the bulk atmosphere (V_a). This reduced the range of the stimulus, V_s, thereby reducing the range of the stomatal response, without indicating low stomatal sensitivity to V. Stomatal responses to V_a may be smaller than expected from V response curves in cuvettes, since V_s rather than the conventionally measured V_a is analogous to V in a well-stirred cuvette. Recently published conclusions that remotely sensed canopy temperatures are inconsistent with stomatal response to V may be based on erroneous estimates of canopy boundary layer conductance and thus of V_s, use of air saturation deficit rather than V to express evaporative demand, and investigation at higher levels of evaporative demand than those eliciting maximal stomatal response.     

塔克拉玛干沙漠南缘豆科与非豆科植物的氮分配

在豆科与非豆科植物光合特性的研究中发现,非豆科植物具有更高的光合速率,与其低的叶氮含量相矛盾。在沙漠中氮素是限制植物生长的关键因子之一,考虑到豆科植物的生物固氮作用和叶氮大部分分配于光合系统,我们假设:(1)非豆科植物具有更低的叶氮含量;(2)分配更少的叶氮于光合系统;(3)具有更高的最大净光合速率(Pmax)和光合氮素利用效率(PNUE)。为了验证这些假设,以塔克拉玛干沙漠南缘的豆科植物骆驼刺(Alhagi sparsifolia)和非豆科植物柽柳(Tamarix ramosissima)、花花柴(Karelinia caspica)为研究对象,比较了它们的叶氮含量、氮分配、Pmax和PNUE等。结果表明:(1)非豆科植物比豆科植物确实有更低的叶氮含量,且差异达到显著水平;(2)非豆科植物分配更少的叶氮于光合系统,但在光合系统内部具有更高效的氮分配机制;(3)非豆科植物具有更高的Pmax和PNUE。在光合系统内部,非豆科植物分配更多的叶氮于羧化系统,而豆科植物分配更多的叶氮于捕光系统。对于非豆科植物而言,其更高的Pmax、PNUE、水分利用效率和表观量子产量,取决于将更多的叶氮投入到羧化和电子传递系统中。这些生理优势决定了塔克拉玛干沙漠南缘非豆科植物高效的资源捕捉和利用能力。     

Allocation of carbon and nitrogen as a function of the internal nitrogen status of a plant: Modelling allocation under non-steady-state situations

In this paper we model allocation of carbon and nitrogen to roots and leaves as a function of the nitrogen status of a plant. Under steady-state conditions, allocation of carbon and nitrogen to leaves is exponentially (positively) correlated with plant nitrogen concentration, whereas allocation to roots is correlated negatively, also in an exponential manner.Allocation functions derived under steady-state conditions are used to simulate biomass partitioning under non-steady-state nutrient conditions. Upon nitrogen deprivation, measured and simulated values are rather similar with time, suggesting that allocation functions derived under steady-state conditions also hold under non-steady-state conditions.     

Photosynthesis,respiration, and carbon allocation of two cool-season perennial grasses in response to surface soil drying

The study was conducted to investigate carbon metabolic responses to surface soil drying for cool-season grasses. Kentucky bluegrass (Poa pratensis L.) and tall fescue (Festuca arundinaceae Schreb.) were grown in a greenhouse in split tubes consisting of two sections. Plants were subjected to three soil moisture regimes: (1) well-watered control; (2) drying of upper 20-cm soil (upper drying); and (3) drying of whole 40-cm soil profile (full drying). Upper drying for 30 d had no dramatic effects on leaf water potential (Ψ_leaf) and canopy photosynthetic rate (P_n) in either grass species compared to the well-watered control, but it reduced canopy respiration rate (R_canopy) and root respiration rate in the top 20 cm of soil (R_top). For both species in the lower 20 cm of wet soil, root respiration rates (R_bottom) were similar to the control levels, and carbon allocation to roots increased with the upper soil drying, particularly for tall fescue. The proportion of roots decreased in the 0-20 cm drying soil, but increased in the lower 20 cm wet soil for both grass species; the increase was greater for tall fescue. The Ψ_leaf, P_n, R_canopy, R_top, R_bottom, and carbon allocation to roots in both soil layers were all significantly higher for upper dried plants than for fully dried plants of both grass species. The reductions in R_canopy and R_top in surface drying soil and increases in root respiration and carbon allocation to roots in lower wet soil could help these grasses cope with surface-soil drought stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Sheathing the blade: Significant contribution of sheaths to daytime and nighttime gas exchange in a grass crop

Despite representing a sizeable fraction of the canopy, very little is known about leaf sheath gas exchange in grasses. Specifically, estimates of sheath stomatal conductance, transpiration and photosynthesis along with their responses to light, CO₂ and vapour pressure deficit (VPD) are unknown. Furthermore, the anatomical basis of these responses is poorly documented. Here, using barley as a model system, and combining leaf-level gas exchange, whole-plant gravimetric measurements, transpiration inhibitors, anatomical observations, and biophysical modelling, we found that sheath and blade stomatal conductance and transpiration were similar, especially at low light, in addition to being genotypically variable. Thanks to high abaxial stomata densities and surface areas nearly half those of the blades, sheaths accounted for up to 17% of the daily whole-plant water use, which -surprisingly- increased to 45% during the nighttime. Sheath photosynthesis was on average 17–25% that of the blade and was associated with lower water use efficiency. Finally, sheaths responded differently to the environment, exhibiting a lack of response to CO₂ but a strong sensitivity to VPD. Overall, these results suggest a key involvement of sheaths in feedback loops between canopy architecture and gas exchange with potentially significant implications on adaptation to current and future climates in grasses.     

A model of the long-term response of carbon allocation and productivity of forests to increased CO2concentration and nitrogen deposition

We present a simple theoretical analysis of the long term response of forest growth and carbon allocation to increased atmospheric [CO₂] and N deposition. Our analysis is based on a recent model which predicts that plant light-use efficiency increases with [CO₂] but is independent of plant N supply. We combine that model with simple assumptions for nitrogen fluxes in the soil. A quasi-equilibrium analysis of the short term tree and soil pools is then used to develop a simple graphical depiction of the long term carbon and nitrogen supply constraints on total growth, stem growth and foliar allocation. Our results suggest that long-term growth responses to [CO₂] and N deposition depend strongly on the extent to which stem allocation and foliage allocation are coupled. At one extreme (‘no coupling’), when stem allocation is fixed and independent of foliage allocation, there is no response of total growth or stem growth to increased [CO₂] unless N deposition increases. At the other extreme (‘linear coupling’), when stem allocation is proportional to foliage allocation, there is a significant long-term increase in total growth following a doubling of [CO₂], even when N deposition is unchanged, but stem growth decreases because of a long-term decrease in foliage allocation. For both types of coupling, total growth and stem growth increase with increasing N deposition. In the case of linear coupling, however, the N deposition response of stem growth is significantly larger than that of total growth, because of a long-term increase in foliage allocation. We compare our results with those obtained previously from an alternative model of canopy light-use efficiency involving a dependence on the foliar N:C ratio in addition to [CO₂]. Our results highlight the need for more experimental information on (i) the extent to which canopy light-use efficiency is independent of N supply, and (ii) the relationship between foliage allocation and stem allocation.     

Precipitation regulates plant gas exchange and its long-term response to climate change in a temperate grassland

Aims Climate change largely impacts ecosystem carbon and water cycles by changing plant gas exchange, which may further cause positive or negative feedback to global climate change. However, long-term global change manipulative experiments are seldom conducted to reveal plant ecophysiological responses to climatic warming and altered precipitation regimes.Methods An 8-year field experiment with both warming and increased precipitation was conducted in a temperate grassland in northern China. We measured leaf gas exchange rates (including plant photosynthesis, transpiration and instantaneous water use efficiency [WUE]) of two dominant plant species (Stipa sareptana var. krylovii and Agropyron cristatum) from 2005 to 2012 (except 2006 and 2010) and those of other six species from 2011 to 2012.Important findings Increased precipitation significantly stimulated plant photosynthetic rates (A) by 29.5% and 19.9% and transpiration rates (E) by 42.2% and 51.2% for both dominant species S. sareptana var. krylovii and A. cristatum, respectively, across the 8 years. Similarly, A and E of the six plant functional types were all stimulated by increased precipitation in 2011 and 2012. As the balance of A and E, the instantaneous WUEs of different plant species had species-specific responses to increased precipitation. In contrast, neither warming nor its interaction with increased precipitation significantly affected plant leaf gas exchange rates. Furthermore, A and E of the two dominant species and their response magnitudes to water treatments positively correlated with rainfall amount in July across years. We did not find any significant difference between the short-term versus long-term responses of plant photosynthesis, suggesting the flexibility of leaf gas exchange under climate change. The results suggest that changing precipitation rather than global warming plays a prominent role in determining production of this grassland in the context of climate change.     

Nitrogen additions and microbial biomass: a meta-analysis of ecosystem studies

Nitrogen (N) enrichment is an element of global change that could influence the growth and abundance of many organisms. In this meta-analysis, I synthesized responses of microbial biomass to N additions in 82 published field studies. I hypothesized that the biomass of fungi, bacteria or the microbial community as a whole would be altered under N additions. I also predicted that changes in biomass would parallel changes in soil CO2 emissions. Microbial biomass declined 15% on average under N fertilization, but fungi and bacteria were not significantly altered in studies that examined each group separately. Moreover, declines in abundance of microbes and fungi were more evident in studies of longer durations and with higher total amounts of N added. In addition, responses of microbial biomass to N fertilization were significantly correlated with responses of soil CO2 emissions. There were no significant effects of biomes, fertilizer types, ambient N deposition rates or methods of measuring biomass. Altogether, these results suggest that N enrichment could reduce microbial biomass in many ecosystems, with corresponding declines in soil CO2 emissions.     

Herbivore-induced changes in plant carbon allocation: assessment of below-ground C fluxes using carbon-14

Effects of above-ground herbivory on short-term plant carbon allocation were studied using maize (Zea mays) and a generalist lubber grasshopper (Romalea guttata). We hypothesized that above-ground herbivory stimulates current net carbon assimilate allocation to below-ground components, such as roots, root exudation and root and soil respiration. Maize plants 24 days old were grazed (c. 25–50% leaf area removed) by caging grasshoppers around individual plants and 18 h later pulse-labelled with¹⁴CO₂. During the next 8 h,¹⁴C assimilates were traced to shoots, roots, root plus soil respiration, root exudates, rhizosphere soil, and bulk soil using carbon-14 techniques. Significant positive relationships were observed between herbivory and carbon allocated to roots, root exudates, and root and soil respiration, and a significant negative relationship between herbivory and carbon allocated to shoots. No relationship was observed between herbivory and¹⁴C recovered from soil. While herbivory increased root and soil respiration, the peak time for¹⁴CO₂ evolved as respiration was not altered, thereby suggesting that herbivory only increases the magnitude of respiration, not patterns of translocation through time. Although there was a trend for lower photosynthetic rates of grazed plants than photosynthetic rates of ungrazed plants, no significant differences were observed among grazed and ungrazed plants. We conclude that above-ground herbivory can increase plant carbon fluxes below ground (roots, root exudates, and rhizosphere respiration), thus increasing resources (e.g., root exudates) available to soil organisms, especially microbial populations.     

Compensatory photosynthesis as a response to partial debudding in ezo spruce, <Emphasis Type="Italic">Picea jezoensis</Emphasis> seedlings

The effects of partial debudding on photosynthesis, stomatal conductance and nitrogen contents of 1-year-old needles and newly grown needles were studied in Picea jezoensis (Sieb. et Zucc.). Seventy-five percent of the buds of P.jezoensis seedlings were removed soon after bud break. Gas exchange was measured three times for 1-year-old needles and twice for current-year needles. The photosynthetic rates of 1-year-old needles were greater in debudded seedlings than in control seedlings, and the difference increased as the growing season progressed. This greater photosynthetic rate in debudded seedlings was accompanied by greater stomatal conductance. However, neither the photosynthetic rates nor the stomatal conductance of current-year needles differed between debudded and control seedlings after the needles had fully expanded. Debudding also had no effect on mass-based nitrogen contents in either the 1-year-old or the current-year needles. Area-based nitrogen in the 1-year-old needles did not differ between debudded and control seedlings, but was greater in debudded seedlings than in control seedlings in current-year needles. These results suggest that the enhanced photosynthetic rate is more likely a result of an increased root/leaf ratio that reduces the stomatal limitation of photosynthetic rate than a result of altered sink-source relationships or increased leaf nitrogen content.     

Assimilation and allocation of carbon and nitrogen of thermal and nonthermal Agrostis species in response to high soil temperature

We studied whether changes in the assimilation and allocation of carbon and nitrogen are associated with plant tolerance to high soil temperatures. Two Agrostis species, thermal Agrostis scabra, a species adapted to high-temperature soils in geothermal areas in Yellowstone National Park (USA), and two cultivars of a cool-season species, Agrostis stolonifera, L-93 and Penncross, were exposed to soil temperatures of 37 or 20 degrees C, while shoots were exposed to 20 degrees C. Net photosynthesis rate, photochemical efficiency, NO(3) (-)-assimilation rate and root viability decreased with increasing soil temperatures in both species. However, the decreases were less pronounced for A. scabra than for both A. stolonifera cultivars. Carbon investment in growth of plants exposed to 37 degrees C decreased more dramatically in both A. stolonifera cultivars than in A. scabra. Nitrogen allocation to shoots was greater in A. scabra than in both creeping bentgrass cultivars at 37 degrees C soil temperature. Our results demonstrate that plant tolerance to high soil temperature is related to efficient expenditure and adjustment of C- and N-allocation patterns between growth and respiration.     

Nitrogen allocation and carbon isotope fractionation in relation to intercepted radiation and position in a young Pinus radiata D. Don tree

The three dimensional distribution of intercepted radiation, intercellular CO₂ concentration (C_i) and late summer needle nitrogen (N) concentration were determined at the tips of all 54 branches in a 6·2-m-tall Pinus radiata D. Don tree growing in a New Zealand plantation. Measurements included above- and below-canopy irradiance, leaf stable carbon isotopic composition (δ¹³C) and tree canopy architecture. The radiation absorption component of the model, MAESTRO, was tested on site and then used to determine the branch tip distribution of intercepted radiation. We hypothesized that in branch tip needles: (i) the allocation of nitrogen and other nutrients would be closely associated with the distribution of intercepted radiation, reflecting carbon gain optimization theory, and (ii) C_i would predominantly reflect changes in photosynthetic rate (A) rather than stomatal conductance (g_s), indicating that the increase in A for a given increase in N concentration was larger than the corresponding increase in g_s. Needle nitrogen concentration was poorly related to intercepted radiation, regardless of the period over which the latter was calculated. At a given height, there was a large azimuthal variation in intercepted radiation but N concentration was remarkably uniform around the tree canopy. There was, however, a linear and positive correspondence between N concentration and δ¹³C and needle height above ground (r² = 0·73 and 0·68, respectively). The very strong linear correspondence between N concentration and C_i (r² = 0·71) was interpreted, using gas exchange measurements, as supporting our second hypothesis. Recognizing the strong apical control in P. radiata and possible effects of leaf nitrogen storage in an evergreen species, we propose that the tree leader must have constituted a very strong carbon sink throughout the growing season, and that the proximity of branch tip needles to the leader affected their photosynthetic capacity and nutrient concentration, independent of intercepted radiation. This implies an integrated internal determination of resource allocation within the tree and challenges the current convention that resources are optimally distributed according to the profile of intercepted radiation.     

Optimal allocation of resources in response to shading and neighbours in the heteroblastic species, Acacia implexa

BACKGROUND AND AIMS: Heteroblasty is an encompassing term referring to ontogenetic changes in the plant shoot. A shaded environment is known to affect the process of heteroblastic development; however, it is not known whether crowded or high density growing conditions can also alter heteroblasty. Compound leaves of the shade-intolerant Acacia implexa allocate less biomass per unit photosynthetic area than transitional leaves or phyllodes and it is hypothesized that this trait will convey an advantage in a crowded environment. Compound leaves also have larger photosynthetic capture area - a trait known to be advantageous in shade. This studied tested the hypothesis that more compound leaves will be developed under shade and crowded environments. Furthermore, this species should undergo optimal allocation of biomass to shoots and roots given shaded and crowded environments. METHODS: A full factorial design of irradiance (high and low) and density levels (high, medium and low) on three populations sourced from varying rainfall regions (high, medium and low) was established under controlled glasshouse conditions. Traits measured include the number of nodes expressing a compound leaf, biomass allocation to shoots and roots, and growth traits. Key Results A higher number of nodes expressed a compound leaf under low irradiance and in high density treatments; however, there were no significant interactions across treatments. Phenotypes strongly associated with the shade avoidance syndrome were developed under low irradiance; however, this was not observed under high density. There was no significant difference in relative growth rates across light treatments, but growth was significantly slower in a crowded environment. Conclusions Heteroblastic development in Acacia can be altered by shade and crowded environments. In this experiment, light was clearly the most limiting factor to growth in a shaded environment; however, in a crowded environment there were additional limiting resources to growth.     

Quantifying the response of photosynthesis to changes in leaf nitrogen content and leaf mass per area in plants grown under atmospheric CO2 enrichment

Previous modelling exercises and conceptual arguments have predicted that a reduction in biochemical capacity for photosynthesis (A_area) at elevated CO₂ may be compensated by an increase in mesophyll tissue growth if the total amount of photosynthetic machinery per unit leaf area is maintained (i.e. morphological upregulation). The model prediction was based on modelling photosynthesis as a function of leaf N per unit leaf area (N_area), where N_area = N_mass×LMA. Here, N_mass is percentage leaf N and is used to estimate biochemical capacity and LMA is leaf mass per unit leaf area and is an index of leaf morphology. To assess the relative importance of changes in biochemical capacity versus leaf morphology we need to control for multiple correlations that are known, or that are likely to exist between CO₂ concentration, N_area, N_mass, LMA and A_area. Although this is impractical experimentally, we can control for these correlations statistically using systems of linear multiple-regression equations. We developed a linear model to partition the response of A_area to elevated CO₂ into components representing the independent and interactive effects of changes in indexes of biochemical capacity, leaf morphology and CO₂ limitation of photosynthesis. The model was fitted to data from three pine and seven deciduous tree species grown in separate chamber-based field experiments. Photosynthetic enhancement at elevated CO₂ due to morphological upregulation was negligible for most species. The response of A_area in these species was dominated by the reduction in CO₂ limitation occurring at higher CO₂ concentration. However, some species displayed a significant reduction in potential photosynthesis at elevated CO₂ due to an increase in LMA that was independent of any changes in N_area. This morphologically based inhibition of A_area combined additively with a reduction in biochemical capacity to significantly offset the direct enhancement of A_area caused by reduced CO₂ limitation in two species. This offset was 100% for Acer rubrum, resulting in no net effect of elevated CO₂ on A_area for this species, and 44% for Betula pendula. This analysis shows that interactions between biochemical and morphological responses to elevated CO₂ can have important effects on photosynthesis.     

Resource allocation to reproduction and soma in Drosophila: a stable isotope analysis of carbon from dietary sugar

Metabolic resources in adults of holometabolous insects may derive either from larval or adult feeding. In Drosophila melanogaster, reproduction and lifespan are differently affected by larval vs. adult resource availability, and it is unknown how larval vs. adult acquired nutrients are differentially allocated to somatic and reproductive function. Here we describe the allocation of carbon derived from dietary sugar in aging female D. melanogaster. Larval and adult flies were fed diets contrasting in sucrose (13)C/(12)C, from which we determined the extent to which carbon acquired at each stage contributed to adult somatic tissue and to egg manufacture. Dietary sugar is very important in egg provisioning; at every age, roughly one half of the carbon in eggs was derived from sugar, which turned over from predominantly larval to entirely adult dietary sources. Sucrose provided approximately 40% of total somatic carbon, of which adult dietary sucrose came to supply approximately 75%. Unlike in eggs, however, adult acquired sucrose did not entirely replace the somatic carbon from larvally acquired sucrose. Because carbon from larval sucrose appears to be fairly "replaceable", larval sucrose cannot be a limiting substrate in resource allocation between reproduction and lifespan.