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     

Reproductive yield of individuals competing for light in a dense stand of an annual, Xanthium canadense

In a dense stand, individuals compete with each other for resources, especially for light. Light availability decreases with increasing depth in the canopy, thus light competition becoming stronger with time in the vegetative phase. In the reproductive phase, on the other hand, leaves start senescing, and the light environment, particularly of smaller individuals, will be improved. To study the effect of change in light climate on reproduction of individuals, we established an experimental stand of an annual, Xanthium canadense, and assessed temporal changes in whole plant photosynthesis through the reproductive phase with particular reference to light availability of individuals. At flowering, 83% of individuals were still alive, but only 27% survived to set seeds. Most of the individuals that died in the reproductive phase were smaller than those that produced seeds. Individuals that died at the early stage of the reproductive phase had a lower leaf to stem mass ratio, suggesting that the fate of individuals was determined partly by the pattern of biomass allocation in this period. At the early stage of the reproductive phase, leaf area index (LAI) of the stand was high and larger individuals had higher whole plant photosynthesis than smaller individuals. Although light availability at later stages was improved with reduction in LAI, whole plant photosynthesis was very low in all individuals due to a lower light use efficiency, which was caused by a decrease in photosynthetic N use efficiency. We conclude that light competition was still strong at the early stage of the reproductive phase and that later improvement of light availability did not ameliorate the photosynthesis of smaller individuals.     

Respiration and reproductive effort in Xanthium canadense

* BACKGROUND AND AIMS: The proportion of resources devoted to reproduction in the plant is called the reproductive effort (RE), which is most commonly expressed as the proportion of reproductive biomass to total plant biomass production (RE(W)). Reproductive yield is the outcome of photosynthates allocated to reproductive structures minus subsequent respiratory consumption for construction and maintenance of reproductive structures. Thus, RE(W) can differ from RE in terms of photosynthates allocated to reproductive structures (RE(P)). * METHODS: Dry mass growth and respiration of vegetative and reproductive organs were measured in Xanthium canadense and the amount of photosynthates and its partitioning to dry mass growth and respiratory consumption were determined. Differences between RE(W) and RE(P) were analysed in terms of growth and maintenance respiration. * KEY RESULTS: The fraction of allocated photosynthates that was consumed by respiration was smaller in the reproductive organ than in the vegetative organs. Consequently, RE(P) was smaller than RE(W). The smaller respiratory consumption in the reproductive organ resulted from its shorter period of existence and a seasonal decline in temperature, as well as a slower rate of maintenance respiration, although the fraction of photosynthates consumed by growth respiration was larger than in the vegetative organs. * CONCLUSIONS: Reproductive effort in terms of photosynthates (RE(P)) was smaller than that in terms of biomass (RE(W)). This difference resulted from respiratory consumption for maintenance, which was far smaller in the reproductive organ than in vegetative organs.     

Leaf lifespan and lifetime carbon balance of individual leaves in a stand of an annual herb, Xanthium canadense

Leaf lifespan in response to resource availability has been documented in many studies, but it still remains uncertain what determines the timing of leaf shedding. Here, we evaluate the lifetime carbon (C) balance of a leaf in a canopy as influenced by nitrogen (N) availability. Stands of Xanthium canadense were established with high-nitrogen (HN) and low-nitrogen (LN) treatments and temporal changes of C gain of individual leaves were investigated with a canopy photosynthesis model. Daily C gain of a leaf was maximal early in its development and subsequently declined. Daily C gain at shedding was nearly zero in HN, while it was still positive in LN. Sensitivity analyses showed that the decline in the daily C gain resulted primarily from the reduction in light level in HN and by the reduction in leaf N in LN. Smaller leaf size in LN than in HN led to higher light levels in the canopy, which helped leaves of the LN stand maintain for a longer period. These results suggest that the mechanism by which leaf lifespan is determined changes depending on the availability of the resource that is most limiting to plant growth.     

Timing of seed germination and the reproductive effort in Xanthium canadense

The effect of different dates of germination on the timing of flowering and the final reproductive yield was examined in a short-day annual plant Xanthium canadense (cocklebur). Delays in germination of 30 and 60 days deferred flower initiation by 2 and 9 days, respectively. Although plants that germinated later were smaller because of the shorter growing period, the reproductive yields did not show as much reduction as the vegetative biomass. The reproductive effort (RE, defined as the ratio of final reproductive yield to the vegetative biomass at the end of the growing season) increased 1.5 and 2.5 times with delays in germination of 30 and 60 days, respectively. A simple model of plant growth was used to analyse the factors involved in the control of RE, which depends only on the dry mass productivity and its partitioning in the reproductive phase, and is independent of the productivity and partitioning in the vegetative phase. Since relative allocation of dry mass to the reproductive part in the reproductive phase was similar for plants with different germination dates, the different REs could be ascribed mainly to differences in productivity of the vegetative parts in the reproductive period. The dependence of RE on plant size is discussed.     

Effects of shift in flowering time on the reproductive output of Xanthium canadense in a seasonal environment

We studied the effects of a change in flowering date on the reproductive output of a short-day annual plant, Xanthium canadense. The flowering date was changed by photoperiodic manipulation to 1 month earlier or later than the natural flowering date. Plants with the natural flowering date attained the highest reproductive output. For those flowering 1 month earlier or later, the reproductive output was decreased by 42% or 23%, respectively. The reproductive output was analyzed as the product of the biomass production during the reproductive period and its allocation to the reproductive organs. Although delay in flowering increased biomass production, it decreased its fractional allocation to the reproductive organs. The highest reproductive output in the natural flowering plants resulted from a compromise between these two effects of flowering. Plants flowering earlier had higher translocation rates to the reproductive organs and accelerated plant senescence. Later flowering caused a reduction in biomass translocation to the reproductive organs and thus extended the reproductive period. These experimental results are discussed in relation to the cost of reproduction and the optimal time for flowering that maximizes the final reproductive output. It is suggested that the natural flowering time maximized the reproductive output while minimizing the cost of reproduction. Received: 11 September 1997 / Accepted: 12 December 1997     

Branching, nitrogen distribution, and carbon gain in solitary plants of an annual herb, Xanthium canadense

The theory of optimal nitrogen (N) distribution predicts that the carbon gain of plants will be maximised when leaves of higher irradiance have higher N content per area (N _area). Most previous studies have examined optimal N distribution without explicitly considering the branching status of plants. I investigated light environment, N distribution and photosynthetic traits of individual leaves of an herbaceous species, Xanthium canadense. X. canadense was grown solitary under high (HN) and low nutrients (LN). Light availability, leaf mass per unit area and N _area were measured for all leaves within plants. Daily photosynthesis of the plants of actual N distribution was compared with those of optimal and constant N distribution. Branch production was facilitated in HN but not in LN plants. N _area was correlated more with leaf order than with leaf light environment. Although N was more limited and the light environment was less heterogeneous within crowns in LN than in HN plants, leaf N distribution was closer to optimal in the latter. These results suggest that leaf N distribution was not optimised in solitary plants of X. canadense. Because this species often regenerates in a dense stand, leaf N distribution might be selected to maximise carbon gain only in such a stand. Leaf N distribution might thus be constrained by the regeneration strategy of the species.     

Nitrogen uptake in a Norway spruce stand following ammonium sulphate application,fertigation, irrigation,drought and nitrogen-free-fertilisation

The above-ground accumulation of N,N uptake and litter quality resulting from improved or deteriorated availability of water and nutrients in a 25 year old Norway spruce stand in SW Sweden (as part of the Skogaby project) is presented. Treatment include irrigation; artificial drought; ammonium sulphate addition; N-free-fertilisation and irrigation with liquid fertilisers including a complete set of nutrients according to the Ingested principle (fertigation). At start of the experiment the stand contained 86.5 t dry mass and 352 kg N ha⁻¹. The following three years the annual N uptake in untreated trees was 32 kg N ha⁻¹ to be compared with the annual N throughfall of 17 kg ha⁻¹. Simultaneously, the treatment with ammonium sulphate and liquid fertilisation resulted in 48 and 56 kg ha⁻¹ y⁻¹, respectively, in treatment specific N-uptake following an application of 100 kg N ha⁻¹ y⁻¹. Addition of a N-free fertiliser resulted in improved N-uptake by 19 kg N ha⁻¹ y⁻¹ and irrigation by 10 kg N ha⁻¹ y⁻¹, compared to control. A linear relation between total above-ground dry mass production and N-uptake was found for trees growing with similar water availability. Dry mass production increased with increased water availability given the same N-uptake. It is concluded that the studied stand this far is not N saturated', as N fertilisation resulted in both increased N uptake and increased growth. Addition of a N-free-fertiliser resulted in increased uptake of N compared to the control, indicating an increased mineralisation rate or uptake capacity of the root system. The linear relation between N uptake and biomass production shows that at this study site N is a highly limiting factor for growth.     

Nitrogen use efficiency in instantaneous and daily photosynthesis of leaves in the canopy of a Solidago altissima stand

Photosynthetic capacity was measured on detached leaves sampled in a canopy of Solidago altissima L. Non-rectangular hyperbola fitted the light response curve of photosynthesis and significant correlations were observed between leaf nitrogen per unit area and four parameters which characterize the light-response curve. Using regressions of the parameters on leaf nitrogen, a model of leaf photosynthesis was constructed which gave the relationships between leaf nitrogen, photon flux density (PFD) and photosynthesis. Curvilinear relations were obtained between leaf nitrogen and photosynthetic rate on both an instantaneous and a daily basis. Nitrogen use efficiency (NUE, photosynthesis per unit leaf nitrogen) was calculated against leaf nitrogen under varying PFDs. The optimum nitrogen content per unit leaf area that maximizes NUE shifted to higher values with increasing PFD. Field measurements of PFD showed high positive correlations between the distribution of leaf nitrogen in the canopy and relative PFD. The predicted optimum leaf nitrogen content for each level in the canopy, to achieve maximized NUE during a clear day, was close to the actual nitrogen distribution as found through sampling.     

A general analysis of resource allocation by competing individuals.

A linear model for population dynamics in a stationary stochastic environment is introduced based on linearizing the N-species Lotka-Volterra competition equations in discrete time. Iteration of the linear model shows the sequence of population sizes to be formed from a simple linear operation on the sequence of carrying capacities. The transfer function for this operation is calculated and the spectral properties of time series data on population size follow directly.The above approach is illustrated with a symmetrical two-species competition system assuming white noise variation in the carrying capacities. The results are interpreted in detail with the following ideas. (1) The intrinsic rate of increase governs the “responsiveness” of the population to changes in the carrying capacity; (2) one effect of competition is to reduce the “effective rate of increase” of the population. Increasing competition can produce effects identical to that of lowering the intrinsic rate of increase; (3) the other effect of competition is to communicate the stochastic variation in one species' carrying capacity to its competitors. The end result of this communication depends critically on the cross-correlation scheme among the carrying capacities of the competing species.     

Zinc uptake by rice,as affected by metabolic inhibitors and competing cations

Summary The effects of zinc application on zinc uptake, distribution and translocation in maize and barley grown in zinc deficient soil with high pH and high calcium content were studied. Zn⁶⁵ content and uptake in roots, sheaths and blades of maize and barley plants increased significantly with increased levels of zinc application. The sheaths contained highest Zn⁶⁵ content followed by roots and blades. The distribution of total zinc, however, differed from that of radioactive zinc. The roots had the highest zinc content, followed by sheaths and blades. The two species differed very little in zinc distribution patterns. The autoradiographs of intact maize and barley plants showed that Zn⁶⁵ was fairly evenly distributed in the main and auxiliary roots, but, there was a relatively higher concentration at the root-stem junction. The Zn⁶⁵ concentration was higher in nodes than in internodes, and in young emerging leaves compared to older leaves. re]19730819 Institute of Soil Science and Isotope Laboratory Agricultural University of Norway     

Nitrogen uptake and use of two contrasting maize hybrids differing in leaf senescence

In eastern Canada, the use of fertilizer N has been identified as the most energy-consuming component of maize (Zea mays L.) grain production. As the economic and environmental costs of excessive N fertilization rise, there is an increased emphasis on selection of hybrids with greater N use efficiency (NUE; defined as the ratio of the amount of ¹⁵N recovered in grain or stover dry matter to the amount of fertilizer ¹⁵N applied to the soil in this study). Using an ¹⁵N-labelling approach, a field study was conducted on a tile-drained Brandon loam soil (Typic Endoaquoll) on the Central Experimental Farm at Ottawa, Canada (45°22 N, 75°43 W) in 1993 and 1994. Fertilizer N uptake and partitioning within the plant in relation to dry matter changes were monitored during development of a current stay-green maize hybrid and an older early-senescing hybrid grown with three fertilizer N levels (0, 100, 200 kg N ha^-1). Dry matter, N concentration and¹⁵ N atom% enrichment of plant components were determined at five growth stages. The current stay-green hybrid, Pioneer 3902 had greater NUE than the old early-senescing hybrid, Pride 5, which was associated with 24% more dry matter production and 20% more N uptake during grain fill for Pioneer 3902. There was no indication of greater allocation of N to the grain in Pioneer 3902. Our data suggest that prolonged maintenance of green leaf area for photosynthate production during grain fill and the ability to take up available soil N later in grain filling are characteristics of maize hybrids with greater NUE.     

陆地生态系统植物的氮源及氮素吸收

氮是植物生长发育所必需的营养元素,也是其主要的限制因子之一.陆地生态系统植物所需氮的来源及植物对氮素的吸收利用均受控于其种类和生长环境.环境条件的改变,一方面可能改变植物生长区原有氮的形态、浓度、赋存方式等,从而改变氮对植物的供给状况;另一方面可能引起植物生长区土壤质量、水分利用状况、光照等的改变,从而产生耦合现象,直接影响植物的生理生态特性,使植物对氮素的吸收利用发生改变,导致植物生长区的种群类型及物种多样性发生改变,并直接影响到生态系统的功能及演替.本文主要对陆地生态系统中高等植物生长发育所需氮素的来源及植物对氮素吸收利用过程中的影响因素进行了综述和讨论,并结合国内外在该领域的研究现状对其研究前景进行了展望.     

Nitrogen uptake of plants affected by windbreaks

Summary The present study was carried out with the windbreaks to determine the effect of wind protection on the uptake of nitrogen by plants. The results of the analysis clearly show that nitrogen uptake is favourably affected by the windbreaks. Nitrogen uptake by the plants in the protected zone at Altona was 52% higher as compared to the unprotected zone. The plants at both the optimal protected and less protected areas at Newton absorbed similar amounts of nitrogen. The increased uptake of nitrogen in the protected zone at Altona was due to the windbreak which favourably influenced the temperature and the evapotranspiration rate. This resulted in the better growth and yield of plants.

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Sommaire La présente étude fut faite à l'aide de brise-vents afin de determiner l'effect de protection du vent en rapport à l'ingestion de l'azote par les plantes. Les résultats de l'analyse montrent clairement que l'ingestion d'azote est favourablement affectée par les brise-vents. L'ingestion d'azote par les plantes de la zone protégée d'Altona fut 52% plus haute que celle de la zone non-protégée. Les plantes, aux endroits de protection maximale et de moindre protection de Newton, ont absorbé des quantités semblables d'azote. L'augmentation de l'ingestion en azote dans la zone protégée d'Altona fut due à l'effect des brisevents qui ont influen?é favourablement la température et le taux d'évapotranspiration. Ceci donna finalement une meilleure croissance et une meilleure productivité des plantes.

     

Nitrogen uptake and preference in a forest understory following invasion by an exotic grass

Plant–soil interactions have been proposed as a causative mechanism explaining how invasive plant species impact ecosystem processes. We evaluate whether an invasive plant influences plant and soil-microbe acquisition of nitrogen to elucidate the mechanistic pathways by which invaders might alter N availability. Using a ¹⁵N tracer, we quantify differences in nitrogen uptake and allocation in communities with and without Microstegium vimineum, a shade-tolerant, C₄ grass that is rapidly invading the understories of eastern US deciduous forests. We further investigate if plants or the microbial biomass exhibit preferences for certain nitrogen forms (glycine, nitrate, and ammonium) to gain insight into nitrogen partitioning in invaded communities. Understory native plants and M. vimineum took up similar amounts of added nitrogen but allocated it differently, with native plants allocating primarily to roots and M. vimineum allocating most nitrogen to shoots. Plant nitrogen uptake was higher in invaded communities due primarily to the increase in understory biomass when M. vimineum was present, but for the microbial biomass, nitrogen uptake did not vary with invasion status. This translated to a significant reduction (P < 0.001) in the ratio of microbial biomass to plant biomass nitrogen uptake, which suggests that, although the demand for nitrogen has intensified, microbes continue to be effective nitrogen competitors. The microbial biomass exhibited a strong preference for ammonium over glycine and nitrate, regardless of invasion status. By comparison, native plants showed no nitrogen preferences and M. vimineum preferred inorganic nitrogen species. We interpret our findings as evidence that invasion by M. vimineum leads to changes in the partitioning of nitrogen above and belowground in forest understories, and to decreases in the microbial biomass, but it does not affect the outcome of plant–microbe–nitrogen interactions, possibly due to functional shifts in the microbial community as a result of invasion.     

Nitrogen uptake by heterotrophic bacteria and phytoplankton in Arctic surface waters

We estimated rates of heterotrophic bacterial and phytoplanktonuptake of nitrate, ammonium, and urea using ¹⁵N-labelled nitrogenand specific metabolic inhibitors of prokaryote and eukaryotenitrogen metabolism in the surface waters of the North Water(northern Baffin Bay) during autumn that were characterizedby the absence of cyanobacteria (comprising prochlorophytes).The percentage of nitrate + ammonium uptake by heterotrophicbacteria ranged between 44 and 78% of the measured total uptakeand was the highest when the phytoplankton biomass was relativelylow (<2 µg Chlorophyll a L^–1). Phytoplanktonaccounted for a larger fraction (e.g., 58–95%) of ureauptake than heterotrophic bacteria. When our results are combinedwith those from previous studies carried out in diverse temperateand polar areas, it appears that heterotrophic bacteria accountfor 25% (14–40%; median and interquartile range) of thetotal nitrate uptake in surface waters with chlorophyll biomass<2 µg L^–1. Estimates of new production computedfrom phytoplankton carbon uptake and f-ratios may be stronglyoverestimated in regions where nitrate uptake by heterotrophicbacteria is high and the biomass of phytoplankton is low.     

Nitrogen uptake and nitrogen use efficiency above and below ground along a topographic gradient of soil nitrogen availability

Nitrogen (N) uptake and nitrogen use efficiency (NUE) are closely related through feedback mechanisms to soil N availability and N cycling in forested ecosystems. We investigated N uptake and NUE not only at the leaf, litterfall, and aboveground levels but also belowground and whole stand levels along a topographic gradient of soil N availability in a cool temperate deciduous forest in Japan. In this study, we addressed how whole stand level N uptake and NUE affect C and N cycling in forested ecosystems. At the leaf, litterfall, and aboveground levels, N uptake decreased and NUE increased with decreasing soil N availability. This pattern resulted from decreasing leaf N concentrations and increasing N resorption efficiencies as soil N availability declined. Low N concentrations in litterfall may have resulted in little soil N being available to plants, due to microbial immobilization. In contrast, when belowground components were included, N uptake and NUE were not correlated with soil N availability. This was mainly due to higher levels of fine root production when soil N availability was low. Higher fine root allocation can result in a high input of detritus to decomposer systems and, thus, contribute to accumulation of soil organic matter and immobilization by microbes, which may result in further soil N availability decline. Our results suggest that allocation to the fine root rather than whole stand level NUE is important for C and N cycling in forested ecosystems, as is the feedback mechanism in which litterfall level NUE shifts with changes in the N concentration of litterfall.     

Nitrogen uptake by ornamental bromeliad: leaf and root efficiency

Plant and Soil - Bromeliads with water-impounding tanks uptake water and nutrients by the leaf-absorbing trichomes present on the leaf surface while the compact root system functions as for plant...     

Nitrogen fixation by white clover when competing with grasses at moderately low temperatures

It was the aim of this study to determine the way in which low temperature modifies the effect of a competing grass on nitrogen fixation of a forage legume. White clover (Trifolium repens L.) was grown in monoculture or in different planting ratios with timothy (Phleum pratense L.) or perennial ryegress (Lolium perenne L.) in growth chambers at either 7.5/5°C (LoT) or 15/10°C (HiT) average day/night temperatures, and with 2.5 or 7.5 mM¹⁵N-labelled nitrate in the nutrient solution.Competition with grass led to a marked increase in the proportion of clover nitrogen derived from symbiosis (% N_sym). This increase was slower at LoT where % N_sym was reduced considerably; it was closely related to the reduction in the amount of available nitrate as a result of its being utilized by the grass.Nitrogen concentration in white clover herbage and dry matter yield per clover plant were reduced, for the most part, when a competing grass was present. The amount of nitrogen fixed per plant of white clover decreased markedly with temperature. Low temperature consequently accentuated competition for nitrate. The capacity of white clover to compete successfully was limited by its slower growth and nitrogen accumulation.