Geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata.

We investigated geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata, to test the hypothesis that escape from herbivory in invasive species permits enhanced growth and lower production of defensive chemicals. We quantified herbivore damage, concentrations of sinigrin, and growth and reproduction inside and outside herbivore exclusion treatments, in field populations in the native and invasive ranges. As predicted, unmanipulated plants in the native range (Hungary, Europe) experienced greater herbivore damage than plants in the introduced range (Massachusetts and Connecticut, USA), providing evidence for enemy release, particularly in the first year of growth. Nevertheless, European populations had consistently larger individuals than US populations (rosettes were, for example, eightfold larger) and also had greater reproductive output, but US plants produced larger seeds at a given plant height. Moreover, flowering plants showed significant differences in concentrations of sinigrin in the invasive versus native range, although the direction of the difference was variable, suggesting the influence of environmental effects. Overall, we observed less herbivory, but not increased growth or decreased defense in the invasive range. Geographical differences in performance and leaf chemistry appear to be due to variation in the environment, which could have masked evolved differences in allocation.

CO(2) enrichment reduces reproductive dominance in competing stands of Ambrosia artemisiifolia (common ragweed).

Plants growing in dense stands may not equally acquire or utilize extra carbon gained in elevated CO(2). As a result, reproductive differences between dominant and subordinate plants may be altered under rising CO(2) conditions. We hypothesized that elevated CO(2) would enhance the reproductive allocation of shaded, subordinate Ambrosia artemisiifolia L. (Asteraceae) individuals more than that of light-saturated dominants. We grew stands of A. artemisiifolia at either 360 or 720 muL L(-1) CO(2) levels and measured the growth and reproductive responses of competing individuals. To test whether elevated CO(2) altered size and reproductive inequalities within stands, we compared stand-level coefficients of variation (CV) in height growth and final shoot, root, and reproductive organ biomasses. Elevated CO(2) enhanced biomass and reduced the CV for all aspects of plant growth, especially reproductive biomass. Allocation to reproduction was higher in the elevated CO(2) than in the ambient treatment, and this difference was more pronounced in small, rather than large plant positive relationships between the CV and total stand productivity declined under elevated CO(2), indicating that growth enhancements to smaller plants diminished the relative biomass advantages of larger plants in increasingly crowded conditions. We conclude that elevated CO(2) stimulates stand-level reproduction while CO(2)-induced growth gains of subordinate A. artemisiifolia plants minimize differences in the reproductive output of small and large plants. Thus, more individuals are likely to produce greater amounts of seeds and pollen in future populations of this allergenic weed.

Phylogenetic overdispersion in Floridian oak communities.

Closely related species that occur together in communities and experience similar environmental conditions are likely to share phenotypic traits because of the process of environmental filtering. At the same time, species that are too similar are unlikely to co-occur because of competitive exclusion. In an effort to explain the coexistence of 17 oak species within forest communities in North Central Florida, we examined correlations between the phylogenetic relatedness of oak species, their degree of co-occurrence within communities and niche overlap across environmental gradients, and their similarity in ecophysiological and life-history traits. We show that the oaks are phylogenetically overdispersed because co-occurring species are more distantly related than expected by chance, and oaks within the same clade show less niche overlap than expected. Hence, communities are more likely to include members of both the red oak and the white + live oak clades than only members of one clade. This pattern of phylogenetic overdispersion arises because traits important for habitat specialization show evolutionary convergence. We hypothesize further that certain conserved traits permit coexistence of distantly related congeners. These results provide an explanation for how oak diversity is maintained at the community level in North Central Florida.

Density-dependent responses of reproductive allocation to elevated atmospheric CO2 in Phytolacca americana.

• This study was conducted to determine whether elevated CO 2 alters patterns of plant reproduction, and whether density affects population- and individual-level responses to elevated CO 2 . • Phytolacca americana was grown in a glasshouse at three population densities under ambient and elevated CO 2 environments, and harvested at both vegetative and seed mature stages. • CO2 did not affect the observed or estimated minimum size required for reproduction. At the population-level, elevated CO2 increased the total and above-ground biomass at both harvests. Density decreased both measurements at the second harvest. At the individual-level, elevated CO2 increased reproductive mass but decreased seed size, and the responses of reproductive allocation were density-dependent. Net photosynthesis at saturating light (Pmax ) increased under elevated CO2 , but decreased with density, with a CO2 × density interaction. • hese results indicate that CO 2 advances timing of flowering by changing growth rate rather than modifying minimum size required for reproduction, while density modifies the responses of reproductive allocations to elevated CO 2 in P. americana .

Tree seedling canopy responses to conflicting photosensory cues.

Light with decreased red:far-red (R:FR) ratios may signal neighbor presence and trigger plant developmental responses. There is some evidence that plant canopies forage towards increased R:FR ratios, but it is unclear to what extent R:FR versus the total amount of photosynthetically active radiation (PAR) influences canopy foraging responses among forest trees. The objective of this study was to examine the relative importance of PAR and R:FR as photosensory cues leading to tree canopy foraging responses. Seedlings of Betula papyrifera Marshall (paper birch) were grown in an experimental garden. Each seedling was germinated and grown in its own shading structure and exposed to two spatially separated light environments, in a factorial design of PAR and R:FR. Plant canopy foraging was evaluated at the end of one growing season in terms of canopy displacement, canopy area, leaf number, direction of stem lean, petiole aspect, and lamina aspect with respect to experimental light treatments. Leaf number and canopy area were greater on the high PAR sides of plants, irrespective of the R:FR treatment. Seedling canopies were displaced towards the direction of high PAR, but this relationship was not significant across all treatments. Petiole aspect was random and showed no significant directedness towards any of the light treatments. Lamina aspect and the direction of stem lean were distributed towards the direction of high PAR, irrespective of the R:FR treatment. Overall, first-year B. papyrifera seedlings used PAR, rather than R:FR ratio, as a photosensory cue for canopy light foraging.

Ion transport by sheep distal airways in a miniature chamber.

Ion transport and the electric profile of distal airways of sheep lungs were studied in a miniature polypropylene chamber with a 1-mm aperture. Small airways with an inner diameter < 1 mm were isolated, opened longitudinally, and then mounted as a flat sheet onto the 1-mm aperture where it was glued and secured with an O-ring. Both sides of the tissue were bathed with identical physiological solutions at 37 degrees C and oxygenated. Pooled data from 27 distal airways showed an inner airway diameter of 854 +/- 22 (SE) microm and a transepithelial potential difference (PD) of 1.86 +/- 0.29 mV, lumen negative. Short-circuit current (I(sc)) was 25 +/- 3.5 microA/cm(2), tissue resistance was 96 +/- 14 Omega, and conductance was 15.2 +/- 1.7 mS/cm(2). At baseline, amiloride-sensitive Na transport accounted for 51% of I(sc) (change in I(sc) = 9.7 +/- 2.6 microA/cm(2); n = 8 airways), corresponding to 0.36 microeq. cm(-2). h(-1). Treatment with 0.1 mM bumetanide did not reduce the I(sc) (n = 5 airways). Exposure to 1 microM Ca ionophore A-23187 raised the I(sc) by 9 microA/cm(2) (47%; P < 0.03; n = 6 airways). The latter effect was blunted by bumetanide. Carbachol at 1 microM provoked a biphasic response, an initial rapid rise in I(sc) followed by a decline (n = 3 airways). There was no significant increase in PD or I(sc) in response to isoproterenol or dibutyryl cAMP. The data suggest that Na absorption constitutes at least 50% of baseline transport activity. Cl or other anion secretion such as HCO(3) appears to be present and could be stimulated by raising intracellular Ca.

Plant biology in the future.

In the beginning of modern plant biology, plant biologists followed a simple model for their science. This model included important branches of plant biology known then. Of course, plants had to be identified and classified first. Thus, there was much work on taxonomy, genetics, and physiology. Ecology and evolution were approached implicitly, rather than explicitly, through paleobotany, taxonomy, morphology, and historical geography. However, the burgeoning explosion of knowledge and great advances in molecular biology, e.g., to the extent that genes for specific traits can be added (or deleted) at will, have created a revolution in the study of plants. Genomics in agriculture has made it possible to address many important issues in crop production by the identification and manipulation of genes in crop plants. The current model of plant study differs from the previous one in that it places greater emphasis on developmental controls and on evolution by differential fitness. In a rapidly changing environment, the current model also explicitly considers the phenotypic variation among individuals on which selection operates. These are calls for the unity of science. In fact, the proponents of "Complexity Theory" think there are common algorithms describing all levels of organization, from atoms all the way to the structure of the universe, and that when these are discovered, the issue of scaling will be greatly simplified! Plant biology must seriously contribute to, among other things, meeting the nutritional needs of the human population. This challenge constitutes a key part of the backdrop against which future evolution will occur. Genetic engineering technologies are and will continue to be an important component of agriculture; however, we must consider the evolutionary implications of these new technologies. Meeting these demands requires drastic changes in the undergraduate curriculum. Students of biology should be trained in molecular, cellular, organismal, and ecosystem biology, including all living organisms.

Interspecific competition in plants: how well do current methods answer fundamental questions?

Accurately quantifying and interpreting the processes and outcomes of competition among plants is essential for evaluating theories of plant community organization and evolution. We argue that many current experimental approaches to quantifying competitive interactions introduce size bias, which may significantly impact the quantitative and qualitative conclusions drawn from studies. Size bias generally arises when estimates of competitive ability are erroneously influenced by the initial size of competing individuals. We employ a series of quantitative thought experiments to demonstrate the potential for size bias in analysis of four traditional experimental designs (pairwise, replacement series, additive series, and response surfaces) either when only final measurements are available or when both initial and final measurements are collected. We distinguish three questions relevant to describing competitive interactions: Which species dominates? Which species gains? and How do species affect each other? The choice of experimental design and measurements greatly influences the scope of inference permitted. Conditions under which the latter two questions can give biased information are tabulated. We outline a new approach to characterizing competition that avoids size bias and that improves the concordance between research question and experimental design. The implications of the choice of size metrics used to quantify both the initial state and the responses of elements in interspecific mixtures are discussed. The relevance of size bias in competition studies with organisms other than plants is also discussed.

Detection of Cl--HCO3- and Na+-H+ exchangers in human airways epithelium.

Molecular species of the Na(+)-H(+) exchanger (NHE) and anion exchanger (AE) gene families and their relative abundance in the human airway regions were assessed utilizing RT-PCR and the RNase protection assay, respectively. Organ donor lung epithelia from various bronchial regions (small, medium, and large bronchi and trachea) were harvested for RNA extraction. Gene-specific primers for the human NHE and AE isoforms were utilized for RT-PCR. Our results demonstrated that NHE1, AE2, and brain AE3 isoforms were expressed in all regions of the human airway, whereas NHE2, NHE3, AE1, and cardiac AE3 were not detected. RNase protection studies for NHE1 and AE2, utilizing glyceraldehyde-3-phosphate dehydrogenase as an internal standard, demonstrated that there were regional differences in the NHE1 mRNA levels in human airways. In contrast, the levels of AE2 mRNA remained unchanged. Differential regional expression of NHE1 isoform may be related to a higher acid load in the tracheal epithelial cells than in epithelia of distal airways. Fluctuations in PCO(2) during inspiration and expiration are probably larger in the tracheal lumen than in the lumen of distal airways with associated larger swings in intracellular pH with each respiratory cycle. Immunohistochemical staining for AE2 protein demonstrated localization to the epithelial cells of human bronchial mucosa.

Consumption rates and food preferences of slugs in a calcareous grassland under current and future CO2 conditions.

This study explored consumption of a generalist herbivore feeding on leaf tissue of various plant species of a calcareous grassland, and tested whether consumption levels and preferences changed when plants were exposed to 5 years of in situ CO2 enrichment. The first part of this experiment tested whether the consumption patterns of slugs (Deroceras reticulatum) observed in single-species feeding tests were altered when slugs were given a choice of food sources. Overall consumption increased 270% when slugs were given a choice, and they preferred having a choice of food sources more than they preferred having any one food source. Surprisingly, slugs consumed fewer legumes and grasses and more non-leguminous forbs when given a choice. In the second part of this experiment, feeding behaviors of slugs in response to elevated CO2 were investigated by feeding them leaves of two legumes, one grass, and a non-leguminous forb (Trifolium medium, Lotus corniculatus, Bromus erectus, and Sanguisorba minor, respectively) in two or four species combinations. In the leguminous species mix, the non-leguminous species mix, and the combined mix (legumes and non-legumes), neither overall consumption by herbivores nor species preference was significantly altered by long-term CO2 enrichment. In the combined species mix, slugs preferred legumes to non-legumes (P=0.012) and exhibited a weak functional group preference shift from non-legumes to legumes (P=0.089) in response to CO2 enrichment. This is the first time such a shift has been observed, and provides evidence that there may be multiple herbivore responses to rising atmospheric CO2 concentrations. Numerous single-species feeding tests using insects have shown that consumption by herbivores may increase when herbivores are fed plants grown in enriched CO2 atmospheres. This study clearly demonstrates the limited applicability of non-choice feeding trials to generalist herbivores in species-rich communities.

Changes in drought response strategies with ontogeny in Quercus rubra: implications for scaling from seedlings to mature trees.

We investigated scaling of physiological parameters between age classes of Quercus rubra by combining in situ field measurements with an experimental approach. In the in situ field study, we investigated changes in drought response with age in seedlings, juveniles, and mature trees of Q. rubra. Throughout the particularly dry summer of 1995 and the unusually wet summer of 1996 in New England, we measured water potential of leaves (ΨLeaf) and gas exchange of plants at three sites at the Harvard Forest in Petersham, Massachusetts. In order to determine what fraction of the measured differences in gas exchange between seedlings and mature trees was due to environment versus ontogeny, an experiment was conducted in which seedlings were grown under light and soil moisture regimes simulating the environment of mature trees. The photosynthetic capacity of mature trees was three-fold greater than that of seedlings during the wet year, and six-fold greater during the drought year. The seedling experiment demonstrated that the difference in photosynthetic capacity between seedlings and mature trees is comprised equally of an environmental component (50%) and an ontogenetic component (50%) in the absence of water limitation. Photosynthesis was depressed more severely in seedlings than in mature trees in the drought year relative to the wet year, while juveniles showed an intermediate response. Throughout the drought, the predawn leaf water potential (ΨPD) of seedlings became increasingly negative (-0.4 to -1.6 MPa), while that of mature trees became only slightly more negative (-0.2 to -0.5 MPa). Again, juveniles showed an intermediate response (-0.25 to -0.8 MPa). During the wet summer of 1996, however, there was no difference in ΨPD between seedlings, juveniles and mature trees. During the dry summer of 1995, seedlings were more responsive to a major rain event than mature trees in terms of ΨLeaf , suggesting that the two age classes depend on different water sources. In all age classes, instantaneous measurements of intrinsic water use efficiency (WUEi), defined as C assimilation rate divided by stomatal conductance, increased as the drought progressed, and all age classes had higher WUEi during the drought year than in the wet year. Mature trees, however, showed a greater ability to increase their WUEi in response to drought. Integrated measurements of WUE from C isotope discrimination (Δ) of leaves indicated higher WUE in mature trees than juveniles and seedlings. Differences between years, however, could not be distinguished, probably due to the strong bias in C isotope fractionation at the time of leaf production, which occurred prior to the onset of drought conditions in 1995. From this study, we arrive at two main conclusions.

Elevated CO2 enhances resprouting of a tropical savanna tree.

The savannas (cerrado) of south-central Brazil are currently subjected to frequent anthropogenic burning, causing widespread reduction in tree density. Increasing concentrations of atmospheric CO2 could reduce the impact of such frequent burning by increasing the availability of nonstructural carbohydrate, which is necessary for resprouting. We tested the hypotheses that elevated CO2 stimulates resprouting and accelerates replenishment of carbohydrate reserves. Using a factorial experiment, seedlings of a common Brazilian savanna tree, Keilmeyera coriacea, were grown at 350 ppm and 700 ppm CO2 and at two nutrient levels. To simulate burning, the plants were either clipped at 15 weeks or were left unclipped. Among unclipped plants, CO2 and nutrients both stimulated growth, with no significant interaction between nutrient and CO2 effects. Among clipped plants, both CO2 and nutrients stimulated resprouting. However, there was a strong interaction between CO2 and nutrient effects, with CO2 having a significant effect only in the presence of high nutrient availability. Under elevated CO2, carbohydrate reserves remained at higher levels following clipping. Root total nonstructural carbohydrate remained above 36% in all treatments, so carbohydrate reserves did not limit regrowth. These results indicate that under elevated CO2 this species may be better able to endure the high frequency of anthropogenic burning in the Brazilian savannas.

Expression of the Na+/H+ and Cl-/HCO-3 exchanger isoforms in proximal and distal human airways.

Recent studies have indicated the presence of Na+/H+ and Cl-/HCO-3 exchange activities in lung alveolar and tracheal tissues of various species. To date, the identity of the Na+/H+ (NHE) and Cl-/HCO-3 (AE) exchanger isoforms and their regional distribution in human airways are not known. Molecular species of the NHE and AE gene families and their relative abundance in the human airway regions were assessed utilizing RT-PCR and the RNase protection assay, respectively. Organ donor lung epithelia from various bronchial regions (small, medium, and large bronchi and trachea) were harvested for RNA extraction. Gene-specific primers for the human NHE and AE isoforms were utilized for RT-PCR. Our results demonstrated that NHE1, AE2, and brain AE3 isoforms were expressed in all regions of the human airways, whereas NHE2, NHE3, AE1, and cardiac AE3 were not detected. RNase protection studies for NHE1 and AE2, utilizing glyceraldehyde-3-phosphate dehydrogenase as an internal standard, demonstrated that there were regional differences in the NHE1 mRNA levels in human airways. In contrast, the levels of AE2 mRNA remained unchanged. Differential expression of these isoforms in the human airways may have functional significance related to the airway absorption and secretion of electrolytes.

Consequences of incongruency in diurnally varying resources for seedlings of Rumex crispus (Polygonaceae).

The incongruency of diurnally varying resources essential to plants may detrimentally affect plants early in their development as indicated by reduced water use efficiency and carbon gain. Typical diurnal patterns of light and CO(2) availability in a midsized temperate herbaceous or forest gap were simulated in specially designed growth chambers. A sinusoidally varying CO(2) treatment (400 ppm minimum, 800 ppm maximum) approximated the diurnal cycle of CO(2) at the soil surface, while a steady-state CO(2) treatment (600 ppm) with the same average CO(2 )concentration provided a control. Crossed with these two CO(2) treatments were two light regimes, one with 3 h of high light (850 µmol·m·s) in the morning (west side of a gap), and the other with 3 h of high light in the afternoon (east side). All treatments received baseline low light (55 µmol·m·s) for 14 h during the day. Rumex crispus was selected as a model species because of its rosette leaves, which grow close to the ground where diurnal CO(2 )variation is greatest. The relative timing of diurnal variations in light and CO(2) significantly affected seedling water use efficiency, carbon gain, and morphology. Total biomass, photosynthetic rates, daily integrated carbon, water use efficiency, and leaf area were enhanced by morning exposure to high light. Seedlings that were exposed to peak values of light and CO(2) incongruently, i.e., those plants receiving intense afternoon light with diurnally varying CO(2), were detrimentally affected relative to control plants receiving intense afternoon light with steady-state CO(2). The results of this experiment indicate that the incongruent availability of required resources-such as light and CO(2)-can detrimentally affect performance relative to when resources are congruent. These contrasting resource regimes can occur on the east and west side of gaps.

Elevated CO2 ameliorates birch response to high temperature and frost stress: implications for modeling climate-induced geographic range shifts.

Despite predictions that both atmospheric CO2 concentrations and air temperature will rise together, very limited data are currently available to assess the possible interactive effects of these two global change factors on temperate forest tree species. Using yellow birch (Betula alleghaniensis) as a model species, we studied how elevated CO2 (800 vs. 400 µl l-1) influences seedling growth and physiological responses to a 5°C increase in summer air temperatures (31/26 vs. 26/21°C day/night), and how both elevated CO2 and air temperature during the growing season influence seedling ability to survive freezing stress during the winter dormant season. Our results show that while increased temperature decreases seedling growth, temperature-induced growth reductions are significantly lower at elevated CO2 concentrations (43% vs. 73%). The amelioration of high-temperature stress was related to CO2-induced reductions in both whole-shoot dark respiration and transpiration. Our results also show that increased summer air temperature, and to a lesser degree CO2 concentration, make dormant winter buds less susceptible to freezing stress. We show the relevance of these results to models used to predict how climate change will influence future forest species distribution and productivity, without considering the direct or interactive effects of CO2.

Phenotypic plasticity and similarity of DNA among genotypes of an annual plant.

When measured directly, rather than inferred from pedigree analyses, the relationship between similarity in phenotype and similarity in DNA sequence was detectable at the level of members of a single population and strongly depended on the environmental context. Genetic divergence among 27 co-occurring genotypes of Abutilon theophrasti, a common annual plant, was less than 5 per cent as revealed by RAPD-PCR analysis based on over 400 bands per genotype. Nevertheless, within this narrow range, there was a positive correlation between genetic similarity and similarity in the performance of genotypes on temperature and moisture gradients, suggesting that plasticity itself has a genetic basis. No relationship was detected, however, when the phenotypic plasticity was expressed in response to gradients of light intensity or soil fertilization, indicating a weaker genetic basis, or suggesting possible involvement of a few genes of major effect.

Effect of elevated CO2 on interactions betwe en the western flower thrips, Frankliniella occidentalis (Thysanoptera: Thripidae) and the common milkweed, Asclepias syriaca.

We measured the effect of elevated CO2 on populations of the western flower thrips, Frankliniella occidentalis and on the amount of leaf damage inflicted by the thrips to one of its host plants, the common milkweed, Asclepias syriaca. Plants grown at elevated CO2 had significantly greater aboveground biomass and C:N ratios, and significantly reduced percentage nitrogen. The number of thrips per plant was not affected by CO2 treatment, but the density of thrips (numbers per gram aboveground biomass), was significantly reduced at high CO2. Consumption by thrips, expressed as the amount of damaged leaf area per capita, was significantly greater at high CO2, and the amount of leaf area damaged by thrips was increased by 33%. However overall leaf area at elevated CO2 increased by 62%, more than compensating for the increase in thrips consumption. The net outcome was that plants at elevated CO2 had 3.6 times more undamaged leaf area available for photosynthesis than plants at ambient CO2, even though they had only 1.6 times the overall amount of leaf area. This study highlights the need for measuring the effects of herbivory at the whole-plant level and also the importance of taking herbivory into account when predicting plant responses to elevated CO2.

Regenerating temperate forest mesocosms in elevated CO2: belowground growth and nitrogen cycling.

The response of temperate forest ecosystems to elevated atmospheric CO2 concentrations is important because these ecosystems represent a significant component of the global carbon cycle. Two important but not well understood processes which elevated CO2 may substantially alter in these systems are regeneration and nitrogen cycling. If elevated CO2 leads to changes in species composition in regenerating forest communities then the structure and function of these ecosystems may be affected. In most temperate forests, nitrogen appears to be a limiting nutrient. If elevated CO2 leads to reductions in nitrogen cycling through increased sequestration of nitrogen in plant biomass or reductions in mineralization rates, long-term forest productivity may be constrained. To study these processes, we established mesocosms of regenerating forest communities in controlled environments maintained at either ambient (375 ppm) or elevated (700 ppm) CO2 concentrations. Mesocosms were constructed from intact monoliths of organic forest soil. We maintained these mesocosms for 2 years without any external inputs of nitrogen and allowed the plants naturally present as seeds and rhizomes to regenerate. We used 15N pool dilution techniques to quantify nitrogen fluxes within the mesocosms at the end of the 2 years. Elevated atmospheric CO2 concentration significantly affected a number of plant and soil processes in the experimental regenerating forest mesocosms. These changes included increases in total plant biomass production, plant C/N ratios, ectomycorrhizal colonization of tree fine roots, changes in tree fine root architecture, and decreases in plant NH4+ uptake rates, gross NH4+ mineralization rates, and gross NH4+ consumption rates. In addition, there was a shift in the relative biomass contribution of the two dominant regenerating tree species; the proportion of total biomass contributed by white birch (Betula papyrifera) decreased and the proportion of total biomass contributed by yellow birch (B. alleghaniensis) increased. However, elevated CO2 had no significant effect on the total amount of nitrogen in plant and soil microbial biomass. In this study we observed a suite of effects due to elevated CO2, some of which could lead to increases in potential long term growth responses to elevated CO2, other to decreases. The reduced plant NH4+ uptake rates we observed are consistent with reduced NH4+ availability due to reduced gross mineralization rates. Reduced NH4+ mineralization rates are consistent with the increases in C/N ratios we observed for leaf and fine root material. Together, these data suggest the positive increases in plant root architectural parameters and mycorrhizal colonization may not be as important as the potential negative effects of reduced nitrogen availability through decreased decomposition rates in a future atmosphere with elevated CO2.

Intra- and inter-specific variation in canopy photosynthesis in a mixed deciduous forest.

Within the same forest, photosynthesis can vary greatly among species and within an individual tree. Quantifying the magnitude of variation in leaf-level photosynthesis in a forest canopy will improve our understanding of and ability to model forest carbon cycling. This information requires extensive sampling of photosynthesis in the canopy. We used a 22-m-tall, four-wheel-drive aerial lift to reach five to ten leaves from the tops of numerous individuals of several species of temperate deciduous trees in central Massachusetts. The goals of this study were to measure light-saturated photosynthesis in co-occurring canopy tree species under field conditions, and to identify sampling schemes appropriate for canopy tree studies with challenging logistics. Photosynthesis differed significantly among species. Even though all leaves measured were canopy-top, sun-acclimated foliage, the more shade-tolerant species tended to have lower light-saturated photosynthetic rates (P max) than the shade-intolerant species. Likewise, leaf mass per area (LMA) and nitrogen content (N) varied significantly between species. With only one exception, the shade-tolerant species tended to have lower nitrogen content on an area basis than the intolerant species, although the LMA did not differ systematically between these ecological types. Light-saturated P max rates and nitrogen content, both calculated on either an area or a mass basis, and the leaf mass to area ratio, significantly differed not only among species, but also among individuals within species (P<0.0001 for both). Differences among species accounted for a greater proportion of variance in the P max rates and the nitrogen content than the differences among individuals within a species (58.5-78.8% of the total variance for the measured parameters was attributed to species-level differences versus 5.5-17.4% of the variance was attributed to differences between individual trees of a given species). Furthermore, more variation is accounted for by differences among leaves in a single individual tree, than by differences among individual trees of a given species (10.7-30.4% versus 5.5-17.4%). This result allows us to compare species-level photosynthesis, even if the sample size of the number of trees is low. This is important because studies of canopy-level photosynthesis are often limited by the difficulty of canopy access. As an alternative to direct canopy access measurements of photosynthesis, it would be useful to find an "easy-to-measure" proxy for light-saturated photosynthetic rates to facilitate modeling forest carbon cycling. Across all species in this study, the strongest correlation was between nitrogen content expressed on an area basis (mmol m-2, N area) and light-saturated P max rate (µmol m-2 s-1, P maxarea) (r 2=0.511). However, within a given species, leaf nitrogen was not tightly correlated with photosynthesis. Our sampling design minimized intra-specific leaf-level variation (i.e., leaves were taken only from the top of the canopy and at only one point in the season). This implies that easy-to-measure trends in nitrogen content of leaves may be used to predict the species-specific light-saturated P max rates.