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A number of studies have shown that under some conditions plants may fully or partially compensate for leaf tissue loss; however, the mechanisms underlying compensatory responses are not well understood. Previous work demonstrated that the annual herb Abutilon theophrasti fully compensated for 75% defoliation, but only when grown in the absence of stem competition. We examined potential mechanisms of compensatory response and how they are influenced by resource limitation (i.e., competition for light). Full compensation for these annual plants was defined as equal final reproductive output in defoliated and control plants. In the current study we observed substantial compensation in defoliated plants growing at low density, despite losing 75% of leaf area prior to the onset of flowering. Plant responses associated with compensation included (1) increased reproductive efficiency, which may in turn may have resulted from increased canopy light penetration and transient increases in leaf-level photosynthetic rates; (2) greater allocation to reproduction (RA); (3) changes in biomass allocation from roots to shoots; (4) lower leaf longevity, and (5) increased percent fruit set. Although some of these responses were also observed in defoliated plants grown at high density, the inability of high-density plants to compensate appeared to result from competition for light; these plants delayed reproduction and continued to produce new leaves late in the growing season after low-density, defoliated plants had shifted allocation of resources to reproduction.

Elevated CO2 may increase dry mass production of canopies directly through increasing net assimilation rate of leaves and also indirectly through increasing leaf area index (LAI). We studied the effects of CO2 elevation on canopy productivity and development in monospecific and mixed (1:1) stands of two co-occurring C3 annual species, Abutilon theophrasti, and Ambrosia artemisiifolia. The stands were established in the glasshouse with two CO2 levels (360 and 700 μl/l) under natural light conditions. The planting density was 100 per m2 and LAI increased up to 2.6 in 53 days of growth. Root competition was excluded by growing each plant in an individual pot. However, interference was apparent in the amount of photons absorbed by the plants and in photon absorption per unit leaf area. Greater photon absorption by Abutilon in the mixed stand was due to different canopy structures: Abutilon distributed leaves in the upper layers in the canopy while Ambrosia distributed leaves more to the lower layers. CO2 elevation did not affect the relative performance and light interception of the two species in mixed stands. Total aboveground dry mass was significantly increased with CO2 elevation, while no significant effects on leaf area development were observed. CO2 elevation increased dry mass production by 30–50%, which was mediated by 35–38% increase in the net assimilation rate (NAR) and 37–60% increase in the nitrogen use efficiency (NUE, net assimilation rate per unit leaf nitrogen). Since there was a strong overall correlation between LAI and aboveground nitrogen and no significant difference was found in the regression of LAI against aboveground nitrogen between the two CO2 levels, we hypothesized that leaf area development was controlled by the amount of nitrogen taken up from the soil. This hypothesis suggests that the increased LAI with CO2 elevation observed by several authors might be due to increased uptake of nitrogen with increased root growth.

Abutilon theophrasti (C3) and Amaranthus retroflexus (C4), were grown from seed at four partial pressures of CO2: 15 Pa (below Pleistocene minimum), 27 Pa (pre-industrial), 35 Pa (current), and 70 Pa (future) in the Duke Phytotron under high light, high nutrient, and wellwatered conditions to evaluate their photosynthetic response to historic and future levels of CO2. Net photosynthesis at growth CO2 partial pressures increased with increasing CO2 for C3 plants, but not C4 plants. Net photosynthesis of Abutilon at 15 Pa CO2 was 70% less than that of plants grown at 35 Pa CO2, due to greater stomatal and biochemical limitations at 15 Pa CO2. Relative stomatal limitation (RSL) of Abutilon at 15 Pa CO2 was nearly 3 times greater than at 35 Pa CO2. A photosynthesis model was used to estimate ribulose-1,5-bisphosphate carboxylase (rubisco) activity (Vcmax), electron transport mediated RuBP regeneration capacity (J max), and phosphate regeneration capacity (PiRC) in Abutilon from net photosynthesis versus intercellular CO2 (A-C i) curves. All three component processes decreased by approximately 25% in Abutilon grown at 15 Pa compared with 35 Pa CO2. Abutilon grown at 15 Pa CO2 had significant reductions in total rubisco activity (25%), rubisco content (30%), activation state (29%), chlorophyll content (39%), N content (32%), and starch content (68%) compared with plants grown at 35 Pa CO2. Greater allocation to rubisco relative to light reaction components and concomitant decreases in J max and PiRC suggest co-regulation of biochemical processes occurred in Abutilon grown at 15 Pa CO2. There were no significant differences in photosynthesis or leaf properties in Abutilon grown at 27 Pa CO2 compared with 35 Pa CO2, suggesting that the rise in CO2 since the beginning of the industrial age has had little effect on the photosynthetic performance of Abutilon. For Amaranthus, limitations of photosynthesis were balanced between stomatal and biochemical factors such that net photosynthesis was similar in all CO2 treatments. Differences in photosynthetic response to growth over a wide range of CO2 partial pressures suggest changes in the relative performance of C3 and C4 annuals as atmospheric CO2 has fluctuated over geologic time.

Previously we examined how limited rooting space and nutrient supply influenced plant growth under elevated atmospheric CO2 concentrations (McConnaughay et al. 1993). We demonstrated that plant growth enhancement under elevated CO2 was influenced more by the concentration of nutrients added to growth containers than to either the total nutrient content per pot or amount or the dimensions of available rooting space. To gain insight into how elevated CO2 atmospheres affect how plants utilize available belowground space when rooting space and nutrient supply are limited we measured the deployment of roots within pots through time. Contrary to aboveground responses, patterns of belowground deployment were most strongly influenced by elevated CO2 in pots of different volume and shape. Further, elevated CO2 conditions interacted differently with limited belowground space for the two species we studied,Abutilon theophrasti, a C3 dicot with a deep taproot, andSetaria faberii, a C4 monocot with a shallow fibrous root system. ForSetaria, elevated CO2 increased the size of the largest region of low root density at the pot surface in larger rooting volumes independent of nutrient content, thereby decreasing their efficiency of deployment. ForAbutilon, plants responded to elevated CO2 concentrations by equalizing the pattern of deployment in all the pots. Nutrient concentration, and not pot size or shape, greatly influenced the density of root growth. Root densities forAbutilon andSetaria were similar to those observed in field conditions, for annual dicots and monocots respectively, suggesting that studies using pots may successfully mimic natural conditions.

Recently, it has been suggested that small pots may reduce or eliminate plant responses to enriched CO2 atmospheres due to root restriction. While smaller pot volumes provide less physical space available for root growth, they also provide less nutrients. Reduced nutrient availability alone may reduce growth enhancement under elevated CO2. To investigate the relative importance of limited physical rooting space separate from and in conjunction with soil nutrients, we grew plants at ambient and double-ambient CO2 levels in growth containers of varied volume, shape, nutrient concentration, and total nutrient content. Two species (Abutilon theophrasti, a C3 dicot with a deep tap root andSetaria faberii, a C4 monocot with a shallow diffuse root system) were selected for their contrasting physiology and root architecture. Shoot demography was determined weekly and biomass was determined after eight and ten weeks of growth. Increasing total nutrients, either by increasing nutrient concentration or by increasing pot size, increased plant growth. Further, increasing pot size while maintaining equal total nutrients per pot resulted in increased total biomass for both species. CO2-induced growth and reproductive yield enhancements were greatest in pots with high nutrient concentrations, regardless of total nutrient content or pot size, and were also mediated by the shape of the pot. CO2-induced growth and reproductive yield enhancements were unaffected by pot size (growth) or were greater in small pots (reproductive yield), regardless of total nutrient content, contrary to predictions based on earlier studies. These results suggest that several aspects of growth conditions within pots may influence the CO2 responses of plants; pot size, pot shape, the concentration and total amount of nutrient additions to pots may lead to over-or underestimates of the CO2 responses of real-world plants.

Three oldfield annual species (Abutilon theophrasti Medic., Ambrosia artemisiifolia L. and Setaria lutescens (Weigel) Hubb.) were investigated. All three developed substantial mycorrhizal infections when inoculated with Glomus etunicatum Becker & Gerd. Mycorrhizal infection dramatically increased phosphorus content and dry weight of both Abutilon and Ambrosia, but did not significantly affect dry weight and only modestly increased phosphorus content of Setaria. These results were consistent with a lower level of infection and much greater root density in Setaria than in the other species. When Abutilon was grown in the presence of Setaria, mycorrhizal infection had no effect on Abutilon phosphorus content or dry weight. The depressive effect of Setaria on the response to inoculation in Abutilon was probably not caused by water soluble allelopathic chemicals from Setaria roots, but soil leachate from Abutilon plants did inhibit infection in other Abutilon plants. The data were consistent with the hypothesis that the very high root density and effective soil exploitation of Setaria reduced the benefit from mycorrhizal infection in Abutilon via phosphorus depletion in a large proportion of the available soil volume. Furthermore, even if mycorrhizal infection were capable of increasing phosphorus content of Abutilon in the presence of Setaria, the very high competitive ability of Setaria for nitrogen in the soil could have reduced the benefit of an enhanced phosphorus content. Carbon isotope ratios were reduced in Abutilon by mycorrhizal infection, indicating a possible reduction in water use efficiency.

One purpose of this study was to determine whether an increase in plant density would result in a decrease in response to mycorrhizal infection (particularly as measured by phosphorus content). Increases in plant density generally result in increases in root density in the volume of soil occupied by the plants. Root density, in turn, largely determines phosphorus uptake. If mycorrhizal plants had significantly higher effective root densities than non-mycorrhizal plants due to the fungal hyphae and thus were more thorough in exploiting a given volume of soil for phosphorus, then a given increase in root density might result in a greater proportional increase in phosphorus uptake for non-mycorrhizal plants than for mycorrhizal plants. Two experiments were performed in which mycorrhizal infection and available soil volume per plant were manipulated; one in which the number of plants within a given pot size was varied (experiment 1), and another in which single plants were grown in pots of differing volume (experiment 2). The two experiments yielded similar results but for apparently different reasons. In the first experiment, for a given increase in root density, non-mycorrhizal plants had a greater proportional increase in phosphorus uptake than mycorrhizal plants. Thus, as predicted, response to mycorrhizal infection was greatest at the lowest planting density (highest available soil volume per plant, lowest root density). In experiment 2, response to infection was also greatest at the highest available soil volume per plant (largest pot), but pot size did not influence root density. These results show that the benefit from mycorrhizal infection may be partly determined by root density and they suggest that plants either occurring in patches of contrasting root density in a given community, or occurring in different communities with inherently different root densities may differ in their reliance upon mycorrhizal fungi for phosphorus uptake.

We examined in the field the effect of the vesicular-arbuscular (VA) mycorhizal symbiosis on the reproductive success of Abutilon theophrasti Medic., an early successional annual member of the Malvaceae. Mycorrhizal infection greatly enhanced vegetative growth, and flower, fruit and seed production, resulting in significantly greater recruitment the following year. In addition, the seeds produced by mycorrhizal plants were significantly larger and contained significantly more phosphorus than seeds from non-mycorrhizal plants, an effect which may improve offspring vigor. Infection by mycorrhizal fungi may thus contribute to the overall fitness of a host plant and strongly influence long-term plant population dynamics.

The response of twenty maternal families of the annual Abutilon theophrasti to two resource gradients, nutrient and light, was investigated. The structure of the population niche for both biomass and reproductive output was found to be quite different on the two gradients. On the light gradient there was a great diversity of responses among the families while on the nutrient gradient the families responded in a similar manner. On both gradients the plants showed a significant genotype/environment interaction. Three strategies for the production of seed variation have been proposed-all offspring are adapted to the same restricted environment, each offspring of an individual is adapted to a particular environment somewhat different thant that of its siblings, and all the offspring are able to grow in a wide range of environments. We found evidence for all three of these strategies amongst the families. The range of responses seen amongst families (of the same species) in this study was as broad as that found in previous studies among species of the old field annual community to which Abutilon theophrasti belongs. This has significant implications to the nature of competitive interactions and to the evolution of differential resource use in plant populations.

Detailed growth analysis in conjunction with information on leaf display and nitrogen uptake was used to interpret competition between Abutilon theophrasti, a C3 annual, and Amaranthus retroflexus, a C4 annual, under ambient (350 μl l-1) and two levels of elevated (500 and 700 μl l-1) CO2. Plants were grown both individually and in competition with each other. Competition caused a reduction in growth in both species, but for different reasons. In Abutilon, decreases in leaf area ratio (LAR) were responsible, whereas decreased unit leaf rate (ULR) was involved in the case of Amaranthus. Mean canopy height was lower in Amaranthus than Abutilon which may explain the low ULR of Amaranthus in competition. The decrease in LAR of Abutilon was associated with an increase in root/shoot ratio implying that Abutilon was limited by competition for below ground resources. The root/shoot ratio of Amaranthus actually decreased with competition, and Amaranthus had a much higher rate of nitrogen uptake per unit of root than did Abutilon. These latter results suggest that Amaranthus was better able to compete for below ground resources than Abutilon. Although the growth of both species was reduced by competition, generally speaking, the growth of Amaranthus was reduced to a greater extent than that of Abutilon. Regression analysis suggests that the success of Abutilon in competition was due to its larger starting capital (seed size) which gave it an early advantage over Amaranthus. Elevated CO2 had a positive effect upon biomass in Amaranthus, and to a lesser extent, Abutilon. These effects were limited to the early part of the experiment in the case of the individually grown plants, however. Only Amaranthus exhibited a significant increase in relative growth rate (RGR). In spite of the transitory effect of CO2 upon size in individually grown plants, level of CO2 did effect final biomass of competitively grown plants. Abutilon grown in competition with Amaranthus had a greater final biomass than Amaranthus at ambient CO2 levels, but this difference disappeared to a large extent at elevated CO2. The high RGR of Amaranthus at elevated CO2 levels allowed it to overcome the difference in initial size between the two species.

Siblings of Abutilon theophrasti, were grown on a nutrient gradient. The plants grown at higher nutrient levels were larger and produced larger and more seeds than plants grown at lower soil nutrient concentrations. There were no differences in germinability of seeds, but the competitive abilities of resulting plants were markedly different. In two different competition experiments designed to eliminate the effects of genotype, seed size, and germination time, by using synchronously germinated seedlings derived from similar size seed from plants grown at different nutrient levels, we found that plants from seeds produced at higher nutrient levels consistently, outperformed plants from seeds produced at the lower nutrient levels. The dominance of seeds produced at higher nutrient levels may be explained by the fact that they had markedly higher concentrations of nitrogen than did seeds produced at lower soil nutrient levels. The additional advantage of increased seed quality to plants controlling more of the nutrient resource than their neighbors would be expected to accelerate their contributions to the gene pool of the population.

Responses to humidity of net photosynthesis and leaf conductance of single attached leaves were examined in populations of herbs from wet soil sites in Beltsville, Maryland and Davis, California, USA. Plants were grown in controlled environments under three conditions which differed in the magnitude of the day-night temperature difference and in daytime air saturation deficit. No population differences in response were found in Abutilon theophrasti. In Amaranthus hybridus stomatal conductance and net photosynthesis were more reduced by increasing leaf to air water vapor pressure difference (VPD) in the population from Beltsville, but only for the growth condition with a constant 25°C temperature. In Chenopodium album, stomatal conductance was more sensitive to VPD in the population from Davis, but only for the growth condition with 28/22°C day/night temperatures. Population differences in the sensitivity to VPD of leaf conductance were associated with differences in leaf area to root weight ratio. The relative reduction of net photosynthesis as VPD increased was greater than, equal to, or less than the relative decrease in substomatal carbon dioxide partial pressure. The pattern depended on species, and on growth condition. From these results one can not conclude that environmental humidity has been a strong selective force in determining sensitivity to humidity of stomatal conductance.