Increasing Plant Size Research Articles

The "stanchezza" (soil sickness) of sweet basil

Seven glasshouse experiments were set up in 1995–1999 to evaluate the efficacy of various treatments in reducing the incidence of “stanchezza” (soil sickness) of sweet basil, an emerging phytopathological problem in the main basil-growing areas of Italy. Complete or partial replacement of the soil was generally effective in increasing plant size and fresh weight of both roots and aerial parts. Steam disinfestation of the soil abolished the symptoms of “stanchezza” during the first growth cycle, but its effect was reduced on the second crop. Leaching the substrate with water and deep tilling had no or only limited effect on plant growth. Amendment with a commercial biostimulating product containing organic phenolic compounds had only partial efficacy. Bench solarization was the most effective means of control of the syndrome and its effect continued for the second crop cycle. No significant reduction of “stanchezza” was achieved with broad-spectrum antibiotic or fungicide mixtures. No significant differences were detected between bacterial and fungal populations in a soil with “stanchezza” and a non-cultivated soil. Further investigation is now required to shed light on the mechanisms underlying this emerging phytopathological problem.

Interactions between atmospheric CO2 concentration and phosphorus nutrition on the formation of proteoid roots in white lupin (Lupinus albus L.)

AbstractAtmospheric [CO2] affects photosynthesis and therefore should affect the supply of carbon to roots. To evaluate interactions between carbon supply and nutrient acquisition, the [CO2] effects on root growth, proteoid root formation and phosphorus (P) uptake capacity were studied in white lupin (Lupinus albus L.) grown hydroponically at 200, 410 and 750 µmol mol−1 CO2, under sufficient (0·25 mm P) and deficient (0·69 µm P) phosphorus. Plant size increased with increasing [CO2] only at high P. Both P deficiency and increasing [CO2] increased the production of proteoid clusters; the increase in response to increased [CO2] was proportionally greater from low to ambient [CO2] than from ambient to high. The activity of phosphoenol pyruvate carboxylase in the proteoid root, the exudation of organic acids from the roots, and the specific uptake of P increased with P deficiency, but were unaffected by [CO2]. Increasing [CO2] from Pleistocene levels to those predicted for the next century increased plant size and allocation to proteoid roots, but did not change the specific P uptake capacity per unit root mass. Hence, rising [CO2] should promote nutrient uptake by allowing lupins to mine greater volumes of soil.

Fluid Milk Processing Costs: Current State and Comparisons

An economic-engineering model is used to derive the theoretically minimum cost of processing and distributing fluid white milk for the state of Maine. This model represents a state-of-the-art milk processing facility and is used to evaluate three questions: 1) the components of total processing costs; 2) whether the cost of milk processing declines with increasing plant size; and 3) the minimum processing volume to financially justify inplant blow-molding technology. The model indicates that significant savings in per-container processing costs can be achieved by increasing plant size. However, distribution costs, related to the geographical distribution of consumer demand and plant location in the state of Maine do not favor large centralized plants.In addition, this model is compared with results published in 1993 to evaluate cost trends over a 7-yr period. The model indicates import shifts to more technologically advanced processing equipment and a dramatic increase in labor costs. Overall, processing costs have risen 2.9% annually above the rate of inflation. Dairies that are unable to respond to increased labor costs through capital investment and expansion will likely find it more difficult to remain competitive in the milk processing industry.

Wind, Size, and Tree Safety

To persist and survive in any environment, a tree must be mechanically reliable and achieve a reasonable factor of safety. This safety factor, which must be adjusted over the course of plant growth, can be computed for stems and roots based on the quotient of the working load and load-bearing capacity of each organ. This method is illustrated for the stems of cherry trees (Prunus serotina) and for the root systems of an arborescent columnar cactus (Pachycereus pringlei) differing in size. The factor of safety of these organs decreases with increasing plant size. The susceptibility of stems and roots to mechanical failure thus increases as plants grow in mass or height. However, the risk of wind-induced tree failure is reduced in the case of cherry trees by the selective wind-failure of small peripheral branches, which reduces the overall drag forces and bending moments acting on trunks. In the case of the columnar cactus, the demand for root water absorption/storage may take precedence over anchorage as plants increase in overall size. These two case studies illustrate that manifold factors of safety must be considered and ranked simultaneously in terms of the probability of damage or death for different environmental risk factors.

Leaf structural and photosynthetic characteristics, and biomass allocation to foliage in relation to foliar nitrogen content and tree size in three Betula species.

Young trees 0.03-1.7 m high of three coexisting Betula species were investigated in four sites of varying soil fertility, but all in full daylight, to separate nutrient and plant size controls on leaf dry mass per unit area (MA), light-saturated foliar photosynthetic electron transport rate (J) and the fraction of plant biomass in foliage (F(L)). Because the site effect was generally non-significant in the analyses of variance with foliar nitrogen content per unit dry mass (N(M)) as a covariate, N(M) was used as an explaining variable of leaf structural and physiological characteristics. Average leaf area (S) and dry mass per leaf scaled positively with N(M) and total tree height (H) in all species. Leaf dry mass per unit area also increased with increasing H, but decreased with increasing N(M), whereas the effects were species-specific. Increases in plant size led to a lower and increases in N(M) to a greater FL and total plant foliar area per unit plant biomass (LAR). Thus, the self-shading probably increased with increasing N(M) and decreased with increasing H. Nevertheless, the whole-plant average M(A), as well as M(A) values of topmost fully exposed leaves, correlated with N(M) and H in a similar manner, indicating that scaling of MA with N(M) and H did not necessarily result from the modified degree of within-plant shading. The rate of photosynthetic electron transport per unit dry mass (J(M)) scaled positively with N(M), but decreased with increasing H and M(A). Thus, increases in M(A) with tree height and decreasing nitrogen content not only resulted in a lower plant foliar area (LAR = F(L)/M(A)), but also led to lower physiological activity of unit foliar biomass. The leaf parameters (J(M), N(M) and M(A)) varied threefold, but the whole-plant characteristic FL varied 20-fold and LAR 30-fold, indicating that the biomass allocation was more plastically adjusted to different plant internal nitrogen contents and to tree height than the foliar variables. Our results demonstrate that: (1) tree height and N(M) may independently control foliar structure and physiology, and have an even greater impact on biomass allocation; and (2) the modified within-plant light availabilities alone do not explain the observed patterns. Although there were interspecific differences with respect to the statistical significance of the relationships, all species generally fit common regressions. However, these differences were consistent, and suggested that more competitive species with inherently larger growth rates also more plastically respond to N and H.

The biomechanics of Pachycereus pringlei root systems

We report on the root system of the large columnar cactus species Pachycereus pringlei to explore the hypothesis that increasing plant size decreases the ability to resist wind-throw but increases the capacity to absorb and store nutrients in roots (i.e., plant size limits the performance of these functions and may shift the performance of one function in favor of another as size increases). Based on 18 plants differing in size, the root system is characterized by a broad and deep bayonet-like root central to a shallow and extensive lateral system of root elements bearing sinker roots near the stem base. All root types have a living secondary cortex and contain wood with a large volume fraction of ray tissues that increases toward the stem base. Wood stiffness and tensile strength are correlated negatively with the ray tissue volume fraction and thus decrease toward the stem base in lateral and bayonet roots. Calculations show that the ability of the bayonet and proximal lateral root elements to resist wind-throw decreases with increasing plant size, whereas the nutrient absorption/storage capacity of the total root system increases with plant size (i.e., a size-dependent shift between these two root functions occurs).

Crown carbon gain and elevated [CO2] responses of understorey saplings with differing allometry and architecture

Abstract1. Attempts at determining the physiological basis of species’ differences, such as the ability to grow in deep shade, have been of limited success. However, this basis is fundamental to predicting species’ responses to rising atmospheric CO2 in the forest understorey. We linked a leaf photosynthesis and a tree architecture model to predict the effects of dynamic and steady state photosynthetic characteristics, crown architecture and elevated atmospheric CO2 concentration ([CO2]) on crown‐level carbon gain (Acrown). Twenty‐four‐h Acrown was modelled for shade‐tolerant Acer rubrum and shade‐intolerant Liriodendron tulipifera saplings growing for three years in a forest understorey under ambient and elevated [CO2] in free‐air CO2 enrichment.2. Two factors best explained Acrown in ambient [CO2]: tree light environment and sapling allometry. Microsite light environment influenced carbon gain via daily photosynthetic photon flux (PFD), average diffuse PFD and sunfleck characteristics. Species differences in specific leaf area (SLA) and size‐related biomass allocation to leaves affected the effective leaf area and hence Acrown.3. At a common above‐ground biomass, small saplings (100 g above‐ground dry mass) of L. tulipifera had higher Acrown than A. rubrum samples due to larger SLA and greater biomass allocation to leaves. Larger saplings of the two species had similar Acrown due to greater carbon allocation to leaves with increasing plant size in A. rubrum vs L. tulipifera. For saplings > 800 g, Acrown was greater in A. rubrum than in L. tulipifera. Enhancement of Acrown by elevated [CO2] on sunny days was similar for both species.4. Overall, though the shade‐tolerant species had lower Acrown than the shade‐intolerant species at a common small size, our results indicate that the relative performance of these species can reverse at larger sizes due to allocational differences. These results suggest that elevated [CO2] may accelerate competition for light between A. rubrum and L. tulipifera as these species grow larger in the understorey.

Size‐Dependent Sex Allocation in a Gynodioecious Wild Strawberry: The Effects of Sex Morph and Inflorescence Architecture

Some models of sex allocation predict that if male fitness gains decelerate faster than female fitness gains with increasing size, then plants should increase allocation to female function with increasing plant size. We tested this prediction in a gynodioecious (females and hermaphrodites) wild strawberry by manipulating plant size in clonally replicated genotypes, by hand‐pollinating with pollen from unrelated donors, and by measuring sex allocation response at two levels: among flowers and within flowers. These methods avoided potential confounding factors that can occur in studies of size‐based allocation under natural conditions, such as genotype‐based variation in size, and size‐based variation in pollen limitation or self‐pollination. They also allow us to determine if inflorescence architecture represented a constraint on the response to plant size. We found that a 75% increase in the vegetative size of hermaphrodites resulted in 78% more flowers, 91% more fruits, 10% more pollen per flower, and 16% more ovules per flower. These responses to plant size represent a significant modification of phenotypic gender at the within‐flower level (pollen:ovule ratio) but not at the among‐flower level (fruit:flower ratio). We also found that while female plants enhanced flower and fruit allocation with plant size to the same degree as hermaphrodites, they increased ovule number per flower to a greater degree. Sex differential effects of plant size such as those just described could influence breeding system evolution if they alter the relative seed fertility of the sex morphs. We also found that inflorescence architecture modified the response to plant size. Specifically, the effect of plant size on ovule number was strongest in basal positions and was absent at the most distal position. This, combined with a trend in the opposite direction for pollen production, resulted in position‐dependent pollen:ovule ratios. Small plants had higher pollen:ovule ratios than large plants at the primary (basal) position, equivalent ratios at intermediate positions, and lower ratios at the quaternary (distal) position. We suggest that this architectural variation in response to plant size may result from structural limitations or physiological trade‐offs within the inflorescence.

Is there evidence for the post‐invasion evolution of increased size among invasive plant species?

Many plant species grow taller and have higher reproductive capacity where they are nonindigenous invaders than where they are native components of the flora. Traditionally, it has been accepted that this is a plastic response to a benign environment, though recently this assumption has been challenged and a genetic basis for increased plant size has been invoked. We tested the hypothesis that the increased size of certain weed species is genetically, rather than environmentally, based. A common environment growth experiment revealed no significant differences in the size of Carduus nutans, Digitalis purpurea, Echium vulgare or Senecio jacobaea sampled from alien (Australia and New Zealand) or native (Britain and continental Europe) habitats. We conclude that post‐invasion genetic changes associated with increased size may be unusual and that the phenomenon, where it occurs, generally reflects a plastic response to a novel environment.

Effect of phalaris (Phalaris aquatica L.) and lucerne (Medicago sativa L.) density on seed yield and regeneration of subterranean clover (Trifolium subterraneum L.)

A mixture of 3 subterranean clover (Trifolium subterraneum L.) cultivars (cvv. Goulburn, Seaton Park, and Dalkeith) was grown with 5–40 plants/m2 of phalaris (Phalaris aquatica L.) or lucerne (Medicago sativa L.) for 3 years at Wagga Wagga, NSW (147°21´E, 35°03´S). Clover seed yield was assessed each year, the number of regenerating clover seedlings was counted, herbage yield of clover and the perennials was measured, and the change in perennial density recorded. The influence of perennials on water availability in spring and light during clover seed set was also examined. The impact of perennial density on clover was assessed using regression analyses. In each of the 3 years, subterranean clover seed yield was negatively related to perennial density, but phalaris suppressed clover seed yield more than equivalent densities of lucerne in 2 of the years. Clover seed yield was positively related (R2 = 0.30–0.85) to clover biomass in spring in all 3 years (except for subterranean clover in phalaris in 1994) and to the proportion of total photosynthetically active radiation reaching the clover canopy beneath the perennial (R2 = 0.33–0.83) in 2 of the 3 years. There was a positive relationship between clover seedling regeneration in autumn and size of the summer seed bank in both years in lucerne (R2 = 0.40–0.76) and in 1 year in phalaris (R2 = 0.76) and a negative association between perennial density and clover seedling regeneration. The rate at which the surface profile (0–37 cm) dried in spring was independent of phalaris and lucerne density over the 3 years and did not differ from that of pure clover. The late season clover cultivar, Goulburn, constituted between 57% and 79% of the seed bank following seed set, substantially more than the earlier flowering cultivars, Seaton Park and Dalkeith. Neither perennial density nor species changed the relative competitiveness of the 3 cultivars. The high seed yield of Goulburn and the lack of a perennial-induced change in surface soil water in spring suggest that growing perennials in association with subterranean clover does not increase the level of moisture stress during clover seed set. The good performance of Goulburn is attributed to superior competitive ability for light in late spring. The basal area occupied by lucerne and phalaris increased with perennial density in each of the first 2 years. However, in the third year, phalaris basal area was similar at all densities. The findings suggest that sowing low densities of lucerne should assist in promoting seed yield and regeneration of subterranean clover. Management of phalaris, which has the capacity to substantially increase plant size at low densities, may require additional strategies such as more frequent grazing in spring.

Allometric scaling of production and life-history variation in vascular plants

A prominent feature of comparative life histories is the well documented negative correlation between growth rate and life span1,2. Patterns of resource allocation during growth and reproduction reflect life-history differences between species1,2. This is particularly striking in tropical forests, where tree species can differ greatly in their rates of growth and ages of maturity but still attain similar canopy sizes3,4. Here we provide a theoretical framework for relating life-history variables to rates of production, dM/dt, where M is above-ground mass and t is time. As metabolic rate limits production as an individual grows, dM/dt ∝ M3/4. Incorporating interspecific variation in resource allocation to wood density, we derive a universal growth law that quantitatively fits data for a large sample of tropical tree species with diverse life histories. Combined with evolutionary life-history theory1, the growth law also predicts several qualitative features of tree demography and reproduction. This framework also provides a general quantitative answer to why relative growth rate (1/M)(dM/df) decreases with increasing plant size (∝M-1/4) and how it varies with differing allocation strategies5,6,7,8.

Economic feasibility of producing ethanol from lignocellulosic feedstocks

A mathematical programming model is built to analyze the economic feasibility of producing ethanol from lignocellulosic feedstocks. The optimal size of an ethanol plant is determined by the trade-off between increasing transportation costs for feedstocks versus decreasing average plant costs as the plant size increases. The ethanol plant is modeled under the assumption that it utilizes recent technological advancements in dilute acid hydrolysis. Potential feedstocks include energy crops, crop residues and woody biomass. It is found that the recent technological advancements appear to make ethanol competitive with gasoline, but only if higher valued chemicals are produced as co-products with the ethanol. The low cost and chemical composition of crop residues make them attractive as a feedstock.

The Bromeliad Microcosm and the Assessment of Faunal Diversity in a Neotropical Forest1

ABSTRACTThe faunas of tank bromeliads were sampled over two years in three forest types at different elevations in the Luquillo Experimental Forest, Puerto Rico, and the diversity of their animal communities compared. Bromeliad plants behaved as islands in that, within forests, the species richness and abundance of their animal communities were significantly and positively correlated with increase in plant size. The amount of canopy debris they accumulated was similarly correlated with increase in plant size. Overall diversity was lowest in the dwarf forest, where plants were uniformly small. Animal communities were stable from year to year, and could be characterised for each forest type and for compartments within the plant. They showed a pattern of high dominance, which increased with elevation (Mc‐Naughton index 37, 54, and 73, respectively, for the tabonuco, palo Colorado, and dwarf forest). Alpha‐diversity for sites sampled in each year reflected net primary productivity (NPP) of the forest, declining with increasing elevation when animal abundance measures were used (jackknife estimates of Simpson's diversity index 6.54 & 11.04 [tabonuco], 3.53 & 6.22 [palo Colorado], and 2.75 & 2.17 [dwarf forest]). Species richness over the two years, however, was highest in the intermediate palo Colorado forest (187 species), compared to 146 and 88 in the tabonuco and dwarf forests, respectively. These figures were close to jackknife estimates of maximum species richness. The difference in species richness between tabonuco and palo Colorado forests was significant in one year only. In addition to NPP, other factors, such as litter quality and the structural complexity of the habitat in the palo Colorado forest, may have influenced species richness. The most abundant species in individual plants were also the most widely occurring, confirming known patterns of abundance and distribution in other functional groups. Diversity within bromeliad microcosms at different elevations supported known relationships between diversity, productivity, and habitat complexity along gradients and was not related to differences in the total bromeliad habitat available for colonization.

540 Field-grown Garden Chrysanthemum Responses to Pinching and Plant Date

Our objective was to determine the effect of planting date and pinching on flowering dates and plant size of field-grown garden mums. Experiments were conducted in the field during two consecutive growing seasons in 1997 and 1998. In one experiment, 15 to 20 cultivars were planted on five dates (14 May, 4 June, 25 June, 16 July, and 4 Aug.) and received no pinching, one manual pinch 2 weeks after potting, or two manual pinches 2 and 4 weeks after potting. In another experiment, four cultivars were planted at the five dates. Pinch treatments were control, one manual pinch, two manual pinches, one Florel spray at 500 mg·L–1, or two Florel sprays at the same time as the manual pinches but on separate plants. Data were collected for days to first color, first open flower, 10 open flowers, and full bloom. Height and width were measured at 10 open blooms. Although the 1998 season was warmer and caused heat delay, the flowering data followed the same trends as the 1997 experiments. Pinching delayed flowering for the early plant dates. Pinching did not affect plant height or plant width. Planting date affected days to 10 blooms for most early season varieties but not late-season varieties. Planting early produced larger plants and more uneven flowering and resulted in greater heat delay of heat-sensitive varieties. Florel delayed flowering and increased plant size. We concluded that pinching was not required to produce high-quality garden mums of many new cultivars.

Impact of saltwater spray andsand deposition on the coastal annualTriplasis purpurea (Poaceae)

Pioneer coastal plants occur in areas where sand movement and airborne salt are common. The objectives of this study were to (1) quantify natural levels of salt and sand deposition in relation to distance from the shoreline of Staten Island, New York, and (2) experimentally determine the impact of saltwater sprays and partial sand burial on growth and reproduction of the native dunegrass Triplasis purpurea. This summer annual matures most seeds in cleistogamous spikelets on leaf sheath-enclosed axillary panicles along culm internodes. Levels of salt deposition onto T. purpurea shoots over 6 d were determined with a conductivity meter to be 175 μg/cm(2) at 39 m from shore in 1997, but declined rapidly with increasing distances to 90 m. Sand deposition over 1 mo in the summer averaged 30 mm at 72-90 m from shore. In a greenhouse factorial experiment, seedlings were unburied, buried to 50% height, or buried to 75% height and simultaneously subjected to no sprays, two sprays/wk, or six sprays/wk of seawater over the summer. Sand deposition increased plant size and seed production, but seawater sprays were mostly detrimental, reducing plant size, seed production, and seed mass. However, T. purpurea tolerated moderate levels of salt deposition. The stimulation of growth and reproduction in partially buried plants is adaptive on the sandy soils. Prolific seed production and tolerance to moderate levels of airborne salt allow this annual to maintain high population densities close to shore.

Structure and Dynamics of Populations of Japanese Barberry (Berberis Thunbergii DC.) in Deciduous Forests of New Jersey

Japanese barberry, Berberis thunbergii DC., has become a prominent exotic species in deciduous forests throughout the eastern and midwestern US. Populations range from small plants occurring at low densities to dense, impenetrable thickets of plants with up to 40 stems/individual. A study was undertaken at Morristown National Historical Park in New Jersey to document plant densities, plant size, recruitment through vegetative processes of new shoot initiation and clonal spread and recruitment from seedling establishment, and mortality of stems and plants. Nearly 2000 shoots on 370 plants were individually marked and followed for two growing seasons, and over 1000 seedlings were also individually marked and followed. Populations vary much more in total shoots/area than they do in plant individuals/area, or in mean plant size (shoots/plant), as even the sparse populations have a few large individuals. Shoot mortality is less than new shoot initiation, but most plants do not change in size or change by small numbers of stems. However, the number of new shoots per plant increases as plant size increases. Once plants have three stems, they suffer little or no mortality. Seedling establishment is proportional to the density of shoots, so that as plants grow in size, local recruitment from seed increases. Large numbers of seedlings, and a survival rate of 10%, combine to make seedling recruitment a major component of population increase. The combination of multiple forms of vegetative and seed-based population growth, and the very low rates of plant mortality due to the multi-stemmed growth form explains the ability of this invasive species to rapidly produce dense, persistent populations.

Seedlings of five boreal tree species differ in acclimation of net photosynthesis to elevated CO(2) and temperature.

Biochemical models of photosynthesis suggest that rising temperatures will increase rates of net carbon dioxide assimilation and enhance plant responses to increasing atmospheric concentrations of CO(2). We tested this hypothesis by evaluating acclimation and ontogenetic drift in net photosynthesis in seedlings of five boreal tree species grown at 370 and 580 &mgr;mol mol(-1) CO(2) in combination with day/night temperatures of 18/12, 21/15, 24/18, 27/21, and 30/24 degrees C. Leaf-area-based rates of net photosynthesis increased between 13 and 36% among species in plants grown and measured in elevated CO(2) compared to ambient CO(2). These CO(2)-induced increases in net photosynthesis were greater for slower-growing Picea mariana (Mill.) B.S.P., Pinus banksiana Lamb., and Larix laricina (Du Roi) K. Koch than for faster-growing Populus tremuloides Michx. and Betula papyrifera Marsh., paralleling longer-term growth differences between CO(2) treatments. Measures at common CO(2) concentrations revealed that net photosynthesis was down-regulated in plants grown at elevated CO(2). In situ leaf gas exchange rates varied minimally across temperature treatments and, contrary to predictions, increasing growth temperatures did not enhance the response of net photosynthesis to elevated CO(2) in four of the five species. Overall, the species exhibited declines in specific leaf area and leaf nitrogen concentration, and increases in total nonstructural carbohydrates in response to CO(2) enrichment. Consequently, the elevated CO(2) treatment enhanced rates of net photosynthesis much more when expressed on a leaf area basis (25%) than when expressed on a leaf mass basis (10%). In all species, rates of leaf net CO(2) exchange exhibited modest declines with increasing plant size through ontogeny. Among the conifers, enhancements of photosynthetic rates in elevated CO(2) were sustained through time across a wide range of plant sizes. In contrast, for Populus tremuloides and B. papyrifera, mass-based photosynthetic rates did not differ between CO(2) treatments. Overall, net photosynthetic rates were highly correlated with relative growth rate as it varied among species and treatment combinations through time. We conclude that interspecific variation may be a more important determinant of photosynthetic response to CO(2) than temperature.

Temperature and ontogeny mediate growth response to elevated CO2 in seedlings of five boreal tree species.

We tested the extent to which growth responses to elevated carbon dioxide (CO2 ) are temperature-dependent and change through early seedling ontogeny among boreal tree species of contrasting relative growth rates (rgr). Populus tremuloides Michx, Betula papyrifera Marsh, Larix laricina (Du Roi) K. Koch, Pinus banksiana Lamb., and Picea mariana (Mill.) B.S.P. were grown from seeds for 3 months in controlled-environment chambers at two CO2 concentrations (370 and 580 μmol mol-1 ) and five temperature regimes of 18/12, 21/15, 24/18, 27/21 and 30/24°C (light/dark). Growth increases in response to CO2 enrichment were minimal at the lowest temperature and maximal at 21/15°C for the three conifers and at 24/18°C or higher for the two broadleaved species, corresponding with differences in optimal temperatures for growth. In both CO2 treatments, rgr among species and temperatures correlated positively with leaf area ratio (lar) (r⩾0·90, P<0·0001). However, at a given lar, rgr was higher in elevated CO2 , owing to enhanced whole-plant net assimilation rate. On average in all species and temperatures at a common plant mass, CO2 enrichment increased rgr (9%) through higher whole-plant net assimilation rate (22%), despite declines in lar in high CO2 (11%). Reductions in lar are thus an important feedback mechanism reducing positive plant growth responses to CO2 . Proportional allocation of dry mass to roots did not vary between CO2 treatments. Early in the experiment, proportional increases in plant dry mass in elevated CO2 were larger in faster-growing Populus tremuloides and B. papyrifera than in the slower-growing conifers. However, growth increases in response to CO2 enrichment fell with time for broadleaved species and increased for the conifers. With increasing plant size over time, compensatory adjustments to CO2 enrichment in the factors that determine rgr, such as lar, were much larger in broadleaves than in conifers. Thus, the hypothesis that faster-growing species are more responsive to elevated CO2 was not supported, given contrasting patterns of growth response to CO2 with increasing plant size and age.

The effect of plant size on the expression of cleistogamy in Mimulus nasutus

1. Most dimorphic angiosperm species produce chasmogamous flowers adapted for allogamy and cleistogamous flowers adapted for autogamy. Plant size is an important internal environmental factor and several studies have suggested an adaptive strategy whereby cleistogamous flowering ensures a basic seed output while larger plants can spare resources to make additional investment in the more expensive but presumably more fit outcrossed seed produced by chasmogamous flowering. This study tests the effect of plant size on the percentage of chasmogamy in a species, Mimulus nasutus, where both the cleistogamous and chasmogamous flowers are autogamous and so seed from chasmogamous flowers has no genetic advantage. 2. We measured change in expression of cleistogamy/chasmogamy in glasshouse‐grown plants of different sizes and architecture. Size was manipulated by varying the length of time for which the plants were grown under short days. Architecture was manipulated by removing stems. 3. We found that increased size led to an increase in the percentage of chasmogamy in M. nasutus. This is unexpected given the lack of genetic advantage of outcrossing and suggests that this effect can occur as a direct result of increased resource availability without the necessity of selection for progeny fitness. 4. An increase in plant size was also found to lead to an increase in resource sinks (developing flowers and seed capsules on other parts of the plant). As increased plant size can therefore produce both increased total resources and increased number of resource sinks, the net resources available to a developing flower will be a balance of these two effects. Resource re‐allocation is shown to occur between flowers on a stem, between stems and between flowers and seed set. In each case, this results in a net decrease in resources to new flowers and so results in there being only sufficient resources for cleistogamous flowering. 5. We discuss the difficulties of using species with varying growth forms to test the hypothesis that expression of allogamous chasmogamy increases with plant size because of the genetic advantage of outcrossing.

Influence of Nodule Development on the Long-term Photosynthetic Response of Beans to Elevated CO2 and Temperature

The widely observed reduction in photosynthetic (Pn) capacity following long-term exposure to elevated CO2 is believed to result from an imbalance in source–sink status. We hypothesized that nitrogen fixation in root nodules would provide a strong sink for photosynthate and lead to a sustained positive photosynthetic response to elevated CO2. Bean plants (Phaseolus vulgaris L., cv Redkloud) were grown in poly chambers at one of four combinations of temperature (35/21 or 26/15°C day/night), and CO2 (350 or 700 ppm). Half the plants in each chamber were inoculated with Rhizobium and fertilized with a complete nutrient solution lacking nitrogen; control plants received a similar solution with nitrogen. Total nitrogenase activity (acetylene reduction assay; 8 weeks after planting) of excised whole root systems was stimulated (up to 4-fold) by elevated CO2, but this response was only significant for 26/15°C-grown plants. Inoculated plants also accumulated more biomass (10%) than control plants. Nodule abundance and size were significantly higher in high CO2-grown plants than ambient CO2 plants, but the Pn capacity of inoculated plants was only slightly greater than that of control plants. Averaged across other treatments, high CO2-grown plants accumulated more biomass (42%) and had higher Pn rates (50%) than ambient CO2 plants. Treatment effects on leaf carbohydrate levels and Pn acclimation to CO2 were not consistent. The results suggest that the higher total nodule activity was due to increased nodule number and size in proportion with increased plant size under high CO2, rather than an increase in nitrogenase activity per nodule. It is also evident that plants with symbiotic nitrogen fixation capability can benefit from elevated CO2, even with reduced input of inorganic nitrogen.