Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N_area) were determined for individuals, and leaf photosynthesis was measured as a function of N_area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P_mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P_mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P_mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P_mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N_area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N_area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.     

藜个体在高密度种群中的氮素利用效率

 氮素利用效率（NUE）是植物养分策略研究中的一项重要内容。该文利用Berendse和Aerts提出的氮素利用效率概念和原理研究了高密度的藜(Chenopodium album)种群中不同植物个体在种内竞争条件下的氮素利用效率。结果表明,由于植株的氮素吸收速率与其个体大小成非线性关系,说明不同植株个体对氮素的竞争属于非对称竞争。个体较大的植株氮素输入较高,而个体较小的植株氮素输出较高,因而较大个体植株的氮素净增加也较高。植株的氮素损失随着个体大小的增加而增加,较大植株个体的氮素浓度随着生长而下降,而较小植株个体的氮素浓度随时间的变化不大,说明个体较小的植株的生长受光照的限制比受氮素的限制更大,而对较大的植株个体而言,它们的生长受氮素的限制更大。高密度藜种群中的不同植物个体具有不同的养分策略,氮素利用效率及其组成部分氮素生产力（NP）和氮素滞留时间（MRT）均不同。植株的NP和MRT与其个体大小正相关,较大的植物个体具有较高的NP和较长的MRT,因而氮素利用效率也高于个体较小的植株。在个体水平上,种内不同植株的NP与MRT不存在权衡关系（Trade-off）。因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分策略,对于研究种内不同植株的养分策略也同样适用。     

向日葵种群中植株个体大小对其氮素利用策略的影响

我们利用Berendse和Aerts提出的氮素利用效率(NUE)概念及原理研究了高密度一年生草本植物向日葵(Helianthus annuus L.)种群中植株个体大小对其氮素吸收利用的影响,并对种内竞争进行了分析.结果表明,植株对氮素的吸收与其个体大小不成线性关系,说明种群内不同植株个体对土壤氮素的竞争属于非对称竞争.植株的氮素损失随着个体大小的增加而增加.个体较大的植株具有较高的氮素输入率和较低的氮素输出率,因而具有较高的氮素净增加值.植株的氮素生产力(NP)和氮素平均滞留时间(MRT)均与植株个体大小呈正相关.较大的植物个体具有较高的NP和较长的MRT,由于NUE为NP和MRT二者的乘积,因而较大个体植株的NUE高于个体较小的植株.同种植物的不同个体的NP和MRT之间不存在协衡关系.氮素回收效率(NRE)与植株个体大小密切相关.在个体水平上,较大的植株个体具有较高的NUE与其较高的NRE有关.种群内植株个体对土壤氮素的非对称竞争主要由于植株对氮素的吸收和利用效率不同所致.因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分利用策略,对于种内不同植株的养分策略研究也同样适用.     

Light acquisition and use by individuals competing in a dense stand of an annual herb, Xanthium canadense

The importance of light acquisition and utilization by individuals in intraspecific competition was evaluated by determining growth and photosynthesis of individual plants in a dense monospecific stand of an annual, Xanthium canadense. Photosynthesis of individual plants in the stand was calculated using a canopy photosynthesis model in which leaf photosynthesis was assumed to be function of leaf nitrogen content and light availability. The estimated photosynthetic rates of individuals were strongly correlated with the measured growth rates. Photosynthetic rates per unit aboveground mass (RPR, relative photosynthetic rate) increased with increasing aboveground mass, suggesting asymmetric (one-sided) competition in the stand. However, larger individuals had similar RPRs, suggesting symmetric (two-sided) competition. These results were consistent with the observation that size inequality over the whole stand increased with growth, but it remained stable among the larger individuals. The RPR of an individual was calculated as the product of absorbed photon flux per unit aboveground mass (Φ_mass) and light use efficiency (LUE, photosynthesis per unit absorbed photon flux). Φ_mass indicates the efficiency of light acquisition, and was higher in larger individuals in the stand, while LUE was highest in individuals with intermediate aboveground mass. LUE depends on leaf nitrogen content. At an early stage, leaf nitrogen contents of smaller individuals were similar to those that maximize LUE. Light availability to smaller individuals decreased as they grew, while their nitrogen contents did not change markedly, which decreased their LUE. We concluded that asymmetric competition among individuals in the stand resulted mainly from lower efficiencies in both light acquisition and light use by smaller individuals. Received: 31 January 1998 / Accepted: 12 November 1998     

向日葵种群中植株个体大小对其氮素利用策略的影响

我们利用Berendse和Aerts提出的氮素利用效率(NUE)概念及原理研究了高密度一年生草本植物向日葵(Helianthus annuus L.)种群中植株个体大小对其氮素吸收利用的影响,并对种内竞争进行了分析。结果表明,植株对氮素的吸收与其个体大小不成线性关系,说明种群内不同植株个体对土壤氮素的竞争属于非对称竞争。植株的氮素损失随着个体大小的增加而增加。个体较大的植株具有较高的氮素输入率和较低的氮素输出率,因而具有较高的氮素净增加值。植株的氮素生产力(NP)和氮素平均滞留时间(MRT)均与植株个体大小呈正相关。较大的植物个体具有较高的NP和较长的MRT,由于NUE为NP和MRT二者的乘积,因而较大个体植株的NUE高于个体较小的植株。同种植物的不同个体的NP和MRT之间不存在协衡关系。氮素回收效率(NRE)与植株个体大小密切相关。在个体水平上,较大的植株个体具有较高的NUE与其较高的NRE有关。种群内植株个体对土壤氮素的非对称竞争主要由于植株对氮素的吸收和利用效率不同所致。因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分利用策略,对于种内不同植株的养分策略研究也同样适用。     

Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure

Wood density (D_t), an excellent predictor of mechanical properties, is typically viewed in relation to support against gravity, wind, snow, and other environmental forces. In contrast, we show the surprising extent to which variation in D_t and wood structure is linked to support against implosion by negative pressure in the xylem pipeline. The more drought-tolerant the plant, the more negative the xylem pressure can become without cavitation, and the greater the internal load on the xylem conduit walls. Accordingly, D_t was correlated with cavitation resistance. This trend was consistent with the maintenance of a safety factor from implosion by negative pressure: conduit wall span (b) and thickness (t) scaled so that (t/b)² was proportional to cavitation resistance as required to avoid wall collapse. Unexpectedly, trends in D_t may be as much or more related to support of the xylem pipeline as to support of the plant.     

Photosynthesis in an invasive grass and native forb at elevated CO2 during an El Niño year in the Mojave Desert

Annual and short-lived perennial plant performance during wet years is important for long-term persistence in the Mojave Desert. Additionally, the effects of elevated CO₂ on desert plants may be relatively greater during years of high resource availability compared to dry years. Therefore, during an El Niño year at the Nevada Desert FACE Facility (a whole-ecosystem CO₂ manipulation), we characterized photosynthetic investment (by assimilation rate-internal CO₂ concentration relationships) and evaluated the seasonal pattern of net photosynthesis (A_net) and stomatal conductance (g_s) for an invasive annual grass, Bromus madritensis ssp. rubens and a native herbaceous perennial, Eriogonum inflatum. Prior to and following flowering, Bromus showed consistent increases in both the maximum rate of carboxylation by Rubisco (V_Cmax) and the light-saturated rate of electron flow (J_max) at elevated CO₂. This resulted in greater A_net at elevated CO₂ throughout most of the life cycle and a decrease in the seasonal decline of maximum midday A_net upon flowering as compared to ambient CO₂. Eriogonum showed significant photosynthetic down-regulation to elevated CO₂ late in the season, but the overall pattern of maximum midday A_net was not altered with respect to phenology. For Eriogonum, this resulted in similar levels of A_net on a leaf area basis as the season progressed between CO₂ treatments, but greater photosynthetic activity over a typical diurnal period. While g_s did not consistently vary with CO₂ in Bromus, it did decrease in Eriogonum at elevated CO₂ throughout much of the season. Since the biomass of both plants increased significantly at elevated CO₂, these patterns of gas exchange highlight the differential mechanisms for increased plant growth. The species-specific interaction between CO₂ and phenology in different growth forms suggests that important plant strategies may be altered by elevated CO₂ in natural settings. These results indicate the importance of evaluating the effects of elevated CO₂ at all life cycle stages to better understand the effects of elevated CO₂ on whole-plant performance in natural ecosystems.     

Photosynthesis, photoinhibition, and nitrogen use efficiency in native and invasive tree ferns in Hawaii

Photosynthetic gas exchange, chlorophyll fluorescence, nitrogen use efficiency, and related leaf traits of native Hawaiian tree ferns in the genus Cibotium were compared with those of the invasive Australian tree fern Sphaeropteris cooperi in an attempt to explain the higher growth rates of S. cooperi in Hawaii. Comparisons were made between mature sporophytes growing in the sun (gap or forest edge) and in shady understories at four sites at three different elevations. The invasive tree fern had 12-13 cm greater height increase per year and approximately 5 times larger total leaf surface area per plant compared to the native tree ferns. The maximum rates of photosynthesis of S. cooperi in the sun and shade were significantly higher than those of the native Cibotium spp (for example, 11.2 and 7.1 µmol m^-2 s^-1, and 5.8 and 3.6 µmol m^-2 s^-1 respectively for the invasive and natives at low elevation). The instantaneous photosynthetic nitrogen use efficiency of the invasive tree fern was significantly higher than that of the native tree ferns, but when integrated over the life span of the frond the differences were not significant. The fronds of the invasive tree fern species had a significantly shorter life span than the native tree ferns (approximately 6 months and 12 months, respectively), and significantly higher nitrogen content per unit leaf mass. The native tree ferns growing in both sun and shade exhibited greater photoinhibition than the invasive tree fern after being experimentally subjected to high light levels. The native tree ferns recovered only 78% of their dark-acclimated quantum yield (F_v/F_m), while the invasive tree fern recovered 90% and 86% of its dark-acclimated F_v/F_m when growing in sun and shade, respectively. Overall, the invasive tree fern appears to be more efficient at capturing and utilizing light than the native Cibotium species, particularly in high-light environments such as those associated with high levels of disturbance.     

Cytochrome oxidase sensitivity to carbon monoxide in leaves of various tall and dwarf wheat cultivars and their crosses

In order to investigate the possible role of Rht genes in the regulation of the redox condition of cytochrome a₃ (cytochrome c oxidase) during steady-state respiration, wheat cultivars belonging to one of two groups - NP 710, NP 846 and NP 875 belonging to the tall group and Olesons dwarf, HD 1982 and HD 2122 of the dwarf group - and the reciprocal crosses between the varieties of these two groups were examined for carbon monoxide (CO) sensitivity in terms of the inhibition of mitochondrial electron transport. Leaves of young wheat seedlings were used. Differences in the redox state of cytochrome a₃ were monitored using the in vivo aerobic assay of nitrate reduction after a 1-min exposure to CO. Dwarf cultivars possessing Rht genes responded marginally (᜖%) to CO inhibition, whereas the response of tall cultivars to CO was higher (51-70%). Since CO forms a complex only with reduced cytochrome a₃, the results indicate differences in the redox state of cytochrome a₃ during in situ respiration of leaves from tall and dwarf plants that are likely to be controlled by cytoplasmic factors.     

Growth and nitrogen use in Xanthium canadense grown in an open or in a dense stand

Plants develop branches profusely when grown solitarily, while less so when grown in a dense stand. Such changes in architecture are associated with changes in dry mass allocation and nitrogen use. Here, we studied what traits in plant growth and nitrogen use were influenced by different light climates in the stand. Annual plants (Xanthium canadense) were grown solitarily or in a dense stand. Dry mass growth was analyzed as the product of the net assimilation rate (NAR) and leaf area (LA). Nitrogen use efficiency (NUE) was analyzed as the product of nitrogen productivity (NP) and the mean residence time (MRT) of nitrogen. These growth variables were further factorized into their components. Solitary plants maintained a high NAR, whereas plants in the dense stand decreased the NAR due to mutual shading. Plants in the dense stand developed a larger LA with a higher specific leaf area than solitary plants. Solitary plants had higher NUE due to higher NP. A temporal increase in NUE was attributed to the increase in MRT of nitrogen. Light climate was different between solitary and dense-stand plants, but they took up a comparable amount of nitrogen and used it differently in response to the given light climate. NUE was thus demonstrated to be a useful tool for analyzing the mechanism leading to different N use in plant growth.     

Dwarf mistletoe affects whole-tree water relations of Douglas fir and western larch primarily through changes in leaf to sapwood ratios

Dwarf mistletoes induce abnormal growth patterns and extreme changes in the biomass allocation of their hosts as well as directly parasitizing them for resources. Because biomass allocation can affect the resource use and efficiency of conifers, we studied the influences of dwarf mistletoe infection on above-ground biomass allocation of Douglas fir and western larch, and the consequences of such changes on whole-tree water use and water relations. Sap flow, tree water potentials, leaf:sapwood area ratios (A_L:A_S), leaf carbon isotope ratios, and nitrogen content were measured on Douglas fir and western larch trees with various degrees of mistletoe infection during the summer of 1996 in western Montana. Heavy dwarf mistletoe infection on Douglas fir and western larch was related to significant increases in A_L:A_S. Correspondingly, water transport dynamics were altered in infected trees, but responses were different for the two species. Higher A_L:A_S ratios in heavily infected Douglas firs were offset by increases in sapwood area-based sap flux densities (Q_SW) such that leaf area-based sap flux densities (Q_L) and predawn leaf water potentials at the end of the summer did not change significantly with mistletoe infection. Small (but statistically insignificant) decreases of Q_L for heavily infected Douglas firs were enough to offset increases in leaf area such that whole-tree water use was similar for uninfected and heavily infected trees. Increased A_L:A_S ratios of heavily infected western larch were not offset by increases of Q_SW. Consequently, Q_L was reduced, which corresponded with significant decreases of water potential at the end of the summer. Furthermore, mistletoe-infection-related changes in A_L:A_S as a function of tree size resulted in greater whole-tree water use for large infected larches than for large uninfected trees. Such changes may result in further depletion of limited soil water resources in mature infected stands late in the growing season. Foliage from infected trees of both species had lower water use efficiencies than non-infected trees. Our results demonstrate substantial changes of whole-tree processes related to mistletoe infection, and stress the importance of integrating whole-tree physiological and structural processes to fully understand the mechanisms by which pathogens suppress forest productivity.     

The effects of elevated CO2 on seed production and seedling recruitment in a sheep-grazed pasture

Seed production and seedling recruitment were measured over 2 years under ambient (360 ppm) and elevated (475 ppm) atmospheric CO₂ in a free air carbon dioxide enrichment (FACE) experiment, carried out in a sheep-grazed pasture on dry, sandy soil in New Zealand. In both years elevated CO₂ led to more dispersed seeds of the grasses Anthoxanthum odoratum, Lolium perenne and Poa pratensis, the legumes Trifolium repens and T. subterraneum and the herbs Hypochaeris radicata and Leontodon saxatilis. The increased seed dispersal in A. odoratum, H. radicata, Leontodon saxatilis and T. repens reflected both more inflorescences per unit area and more seeds per inflorescence under elevated CO₂. The increased seed dispersal in Lolium perenne, P. pratensis and T. subterraneum was due solely to more inflorescences per unit area. The number of seedlings that emerged and survived to at least 7 months of age was increased by elevated CO₂ for H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in both years and for A. odoratum and Lolium perenne in the first year. For species where increased seedling recruitment was noted, there was a significant positive correlation between seed production in summer and seedling emergence in the following autumn and winter, and sowing 200 extra seeds per species m^-2 resulted in more seedlings compared to unsown controls. Elevated CO₂ did not affect seedling survival in any species. There was no measurable effect of elevated CO₂ on canopy and soil surface conditions or soil moisture at the time of seedling emergence. The results suggest the dominant effect of elevated CO₂ on seedling recruitment in this pasture was an indirect one, reflecting effects on the number of seeds produced. The biomass of H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in the above-ground vegetation was greater under elevated than ambient CO₂. However, the size of individual seedlings and mature plants of these four species was unaffected by elevated CO₂. The results indicate an important way elevated CO₂ influenced plant species composition in this pasture was through changes in the pattern of seedling recruitment.     

Effects of elevated CO<Subscript>2</Subscript> on foliar chemistry of saplings of nine species of tropical tree

This study examined the effects of elevated CO₂ on secondary metabolites for saplings of tropical trees. In the first experiment, nine species of trees were grown in the ground in open-top chambers in central Panama at ambient and elevated CO₂ (about twice ambient). On average, leaf phenolic contents were 48% higher under elevated CO₂. Biomass accumulation was not affected by CO₂, but starch, total non-structural carbohydrates and C/N ratios all increased. In a second experiment with Ficus, an early successional species, and Virola, a late successional species, treatments were enriched for both CO₂ and nutrients. For both species, nutrient fertilization increased plant growth and decreased leaf carbohydrates, C/N ratios and phenolic contents, as predicted by the carbon/nutrient balance hypothesis. Changes in leaf C/N levels were correlated with changes in phenolic contents for Virola (r=0.95, P<0.05), but not for Ficus. Thus, elevated CO₂, particularly under conditions of low soil fertility, significantly increased phenolic content as well as the C/N ratio of leaves. The magnitude of the changes is sufficient to negatively affect herbivore growth, survival and fecundity, which should have impacts on plant/herbivore interactions.     

Water availability and carbon isotope discrimination in conifers

The stable C isotope composition ('¹³C) of leaf and wood tissue has been used as an index of water availability at both the species and landscape level. However, the generality of this relationship across species has received little attention. We compiled literature data for a range of conifers and examined relationships among landscape and environmental variables (altitude, precipitation, evaporation) and '¹³C. A significant component of the variation in '¹³C was related to altitude (discrimination decreased with altitude in stemwood, 2.53‰ km^-1 altitude, r²=0.49, and in foliage, 1.91‰ km^-1, r²=0.42), as has been noted previously. The decrease in discrimination with altitude was such that the gradient in CO₂ partial pressure into the leaf (P_a-P_i) and altitude were generally unrelated. The ratio of precipitation to evaporation (P/E) explained significant variation in P_a-P_i of stemwood (r²=0.45) and foliage (r²=0.27), but only at low (<0.8) P/E. At greater P/E there was little or no relationship, and other influences on '¹³C probably dominated the effect of water availability. We also examined the relationship between plant drought stress (O) and '¹³C within annual rings of stemwood from Pinus radiata and Pinus pinaster in south-western Australia. Differential thinning and fertiliser application produced large differences in the availability of water, nutrients and light to individual trees. At a density of 750 stems ha^-1, O and '¹³C were less (more negative) than at 250 stems ha^-1 indicating greater drought stress and less efficient water use, contrary to what was expected in light of the general relationship between discrimination and P/E. The greater '¹³C of trees from heavily thinned plots may well be related to an increased interception of radiation by individual trees and greater concentrations of nutrients in foliage - attributes that increase rates of photosynthesis, reduce P_i and increase '¹³C. '¹³C was thus modified to a greater extent by interception of radiation and by nutrient concentrations than by water availability and the '¹³C-O relationship varied between thinning treatments. Within treatments, the relationship between '¹³C and O was strong (0.38<r²<0.58). We conclude that '¹³C may well be a useful indicator of water availability or drought stress, but only in seasonally dry climates (P/E<1) and where variation in other environmental factors can be accounted for.     

CO2 exchange and thallus nitrogen across 75 contrasting lichen associations from different climate zones

Aiming to investigate whether a carbon-to-nitrogen equilibrium model describes resource allocation in lichens, net photosynthesis (NP), respiration (R), concentrations of nitrogen (N), chlorophyll (Chl), chitin and ergosterol were investigated in 75 different lichen associations collected in Antarctica, Arctic Canada, boreal Sweden, and temperate/subtropical forests of Tenerife, South Africa and Japan. The lichens had various morphologies and represented seven photobiont and 41 mycobiont genera. Chl a, chitin and ergosterol were used as indirect markers of photobiont activity, fungal biomass and fungal respiration, respectively. The lichens were divided into three groups according to photobiont: (1) species with green algae, (2) species with cyanobacteria, and (3) tripartite species with green algal photobionts and cyanobacteria in cephalodia. Across species, thallus N concentration ranged from 1 to 50 mg g^-1 dry wt., NP varied 50-fold, and R 10-fold. In average, green algal lichens had the lowest, cyanobacterial Nostoc lichens the highest and tripartite lichens intermediate N concentrations. All three markers increased with thallus N concentration, and lichens with the highest Chl a and N concentrations had the highest rates of both P and R. Chl a alone accounted for ca. 30% of variation in NP and R across species. On average, the photosynthetic efficiency quotient [K_F=(NP_max+R)/R)] ranged from 2.4 to 8.6, being higher in fruticose green algal lichens than in foliose Nostoc lichens. The former group invested more N in Chl a and this trait increased NP_max while decreasing R. In general terms, the investigated lichens invested N resources such that their maximal C input capacity matched their respiratory C demand around a similar (positive) equilibrium across species. However, it is not clear how this apparent optimisation of resource use is regulated in these symbiotic organisms.     

Feeding rates of the woodlouse Armadillidium vulgare on herb litters produced at two levels of atmospheric CO2

The consumption and assimilation rates of the woodlouse Armadillidium vulgare were measured on leaf litters from five herb species grown and naturally senesced at 350 and 700 µl l^-1 CO₂. Each type of litter was tested separately after 12, 30 and 45 days of decomposition at 18°C. The effects of elevated CO₂ differed depending on the plant species. In Medicago minima (Fabaceae), the CO₂ treatment had no significant effect on consumption and assimilation. In Tyrimnus leucographus (Asteraceae), the CO₂ treatment had no significant effect on consumption, but the elevated CO₂ litter was assimilated at a lower rate than the ambient CO₂ litter after 30 days of decomposition. In the three other species, Galactites tomentosa (Asteraceae), Trifolium angustifolium (Fabaceae) and Lolium rigidum (Poaceae), the elevated CO₂ litter was consumed and/or assimilated at a higher rate than the ambient CO₂ litter. Examination of the nitrogen contents in these three species of litter did not support the hypothesis of compensatory feeding, i.e. an increase in woodlouse consumption to compensate for low nitrogen content of the food. Rather, the results suggest that in herbs that were unpalatable at the start of the experiment (Galactites, Trifolium and Lolium), more of the the litter produced at 700 µl l^-1 CO₂ was consumed than of that produced at 350 µl l^-1 because inhibitory factors were eliminated faster during decomposition.     

Tree seedling growth in natural deep shade: functional traits related to interspecific variation in response to elevated CO2

The mechanisms for species-specific growth responses to changes in atmospheric CO₂ concentration within narrow ecological groups of species, such as shade-tolerant, late-successional trees, have rarely been addressed and are not well understood. In this study the underlying functional traits for interspecific variation in the biomass response to elevated CO₂ were explored for seedlings of five late-successional temperate forest tree species (Fagus sylvatica, Acer pseudoplatanus, Quercus robur, Taxus baccata, Abies alba). The seedlings were grown in the natural forest understorey in very low and low light microsites (an average of 1.3% and 3.4% full sun in this experiment), and were exposed to either current ambient CO₂ concentrations, 500, or 660 µl CO₂ l^-1 in 36 open-top chambers (OTC) over two growing seasons. Even across the narrow range of successional status and shade tolerance, the study species varied greatly in photosynthesis, light compensation point, leaf dark respiration (R_d), leaf nitrogen concentration, specific leaf area (SLA), leaf area ratio (LAR), and biomass allocation among different plant parts, and showed distinct responses to CO₂ in these traits. No single species combined all characteristics traditionally considered as adaptive to low light conditions. At very low light, the CO₂ stimulation of seedling biomass was related to increased LAR and decreased R_d, responses that were observed only in Fagus and Taxus. At slightly higher light levels, interspecific differences in the biomass response to elevated CO₂ were reversed and correlated best with leaf photosynthesis. The data provided here contribute to a mechanistic process-based understanding of distinct response patterns in co-occurring tree species to elevated CO₂ in natural deep shade. I conclude that the high variation in physiological and morphological traits among late-successional species, and the consequences for their responses to slight changes in resource availability, have previously been underestimated. The commonly used broad definitions of functional groups of species may not be sufficient for the understanding of recruitment success and dynamic changes in species composition of old-growth forests in response to rising concentrations of atmospheric CO₂.     

Effect of Nitrogen Supply on the Nitrogen Use Efficiency of an Annual Herb, Helianthus annuus L.

Nitrogen use efficiency (NUE) is the product of nitrogen productivity (NP) and the mean residence time of nitrogen (MRT). Theory suggests that there should be a trade-off between both components,but direct experimental evidence is still scarce. To test this hypothesis, we analyzed the effect of varying nitrogen supply levels on NUEand its two components (NP, MRT) in Helianthus annuus L., an annual herb.The plants investigated were subjected to six nitrogen levels (0, 2, 4, 8, 16, and 32 g N/m2). Total plant production increased substantially with increasing nitrogen supply. Nitrogen uptake and loss also in creased with nitrogen supply. Nitrogen influx (rin) and outflux (rout) were defined as the rates of nitrogen uptake and loss per unit aboveground nitrogen, respectively. Both rin and rout increased with increasing nitrogen supply. In addition, rin was far higher than rout. Consequently, the relative rate of nitrogen incre ment (rin- rout) also increased with nitrogen supply. There were marked differences between treatments with respect to parameters related to the stress resistance syndrome: nitrogen pool size, leaf nitrogen concentration,and net aboveground productivity increased with nitrogen supply. Plants at high nitrogen levels showed a higher NP (the growth rate per unit aboveground nitrogen) and a shorter MRT (the inverse of rout), whereas plants at low nitrogen levels displayed the reverse pattern. Shorter MRT for plants at high nitrogen levels was caused by the abscission of leaves that contained relatively large fractions of total plant nitrogen. We found a negative relationship between NP and MRT, the components of NUE, along the gradient of nitrogen availability, suggesting that there was a trade-off between NP and MRT. The NUE increased with increasing nitrogen availability, up to a certain level, and then decreased. These results offer support for the hypoth esis that adaptation to infertile habitats involves a low nitrogen loss (long MRT in the plant) rather than a high NUE per se. The higher NUE at the plant level was a result, in part, of greater nitrogen resorption during senescence. We suggest that a long MRT (an index of nitrogen conservation) is a potentially successful strategy in nitrogen-poor environments.     

Limitation of stomatal conductance by hydraulic traits: sensing or preventing xylem cavitation?

We tested the hypothesis that hydraulic conductance per unit leaf surface area of plant shoots (K_SL) determines the maximum diurnal stomatal conductance (g_L) that can be reached by plants growing in the field. A second hypothesis was tested that some xylem cavitation cannot be avoided by transpiring plants and might act as a signal for regulating g_L. Eleven woody species were studied, differing from each other with respect to taxonomy, wood anatomy and leaf habit. Maximum diurnal g_L, transpiration rate (E_L), pre-dawn and minimum diurnal leaf water potential (O_pd and O_min, respectively) were measured in the field. The critical O level at which stem cavitation was triggered (O_cav) was measured on detached branches, using the acoustic method. A high-pressure flow meter was used to measure maximum K_SL of 1-year-old shoots. Both g_L and E_L were positively related to K_SL. The whole-plant hydraulic conductance per unit leaf area (K_WL) of all the species studied, calculated as the ratio of E_L to (O (=O_pd-O_min) was closely related to K_SL. In every case, O_min (ranging between -0.85 and -1.35 MPa in the different species) dropped to the O_cav range or was <O_cav (ranging between -0.71 and -1.23 MPa), thus suggesting that some cavitation-induced embolism could not be avoided. The possibility is discussed that some cavitation-induced reduction in K_SL is the signal for stomatal closure preventing runaway embolism. The lack of correlation of g_L to O_cav is discussed in terms of the inconsistency of O_cav as an indicator of the vulnerability of plants to cavitation. No differences in hydraulic traits were observed between evergreen and deciduous species.     

The effect of elevated carbon dioxide and ozone on leaf- and branch-level photosynthesis and potential plant-level carbon gain in aspen

Two aspen (Populus tremuloides Michx.) clones, differing in O₃ tolerance, were grown in a free-air CO₂ enrichment (FACE) facility near Rhinelander, Wisconsin, and exposed to ambient air, elevated CO₂, elevated O₃ and elevated CO₂+O₃. Leaf instantaneous light-saturated photosynthesis (P_S) and leaf areas (A) were measured for all leaves of the current terminal, upper (current year) and the current-year increment of lower (1-year-old) lateral branches. An average, representative branch was chosen from each branch class. In addition, the average photosynthetic rate was estimated for the short-shoot leaves. A summing approach was used to estimate potential whole-plant C gain. The results of this method indicated that treatment differences were more pronounced at the plant- than at the leaf- or branch-level, because minor effects within modules accrued in scaling to plant level. The whole-plant response in C gain was determined by the counteracting changes in P_S and A. For example, in the O₃-sensitive clone (259), inhibition of P_S in elevated O₃ (at both ambient and elevated CO₂) was partially ameliorated by an increase in total A. For the O₃-tolerant clone (216), on the other hand, stimulation of photosynthetic rates in elevated CO₂ was nullified by decreased total A.