Influence of temporal heterogeneity in nitrogen supply on competitive interactions in a desert shrub community

Soil nutrients in arid systems are supplied to plants in brief pulses following precipitation inputs. While these resource dynamics have been well documented, little is known about how this temporal heterogeneity influences competitive interactions. We examined the impacts of the temporal pattern of N supply on competitive intensity and ability in an N-limited desert shrub community. At our field site, the three codominant shrubs, Atriplex confertifolia, A. parryi, and Sarcobatus vermiculatus, differ in seasonal growth patterns, with A. confertifolia and S. vermiculatus achieving higher growth rates earlier in the growing season than A. parryi. We predicted that these timing differences in maximum growth rate may interact with temporal variation in N supply to alter competitive abilities over time. Seedlings of the two Atriplex species were planted either individually in field plots or as target plants surrounded by neighbor seedlings. After one year of establishment, the same amount of ¹⁵N was applied to plots either as early spring pulses, mid spring pulses or continuously through the second growing season. Competitive effects were observed under continuous and pulsed N supply. Averaged across all target–neighbor treatments, competitive intensity was ∼1.8-fold greater when N was pulsed compared to when N was supplied continuously, but overall, the outcome of competitive interactions was not influenced by N pulse timing. While the timing of resource supply did not differentially influence the competitive abilities of coexisting species in this system, the temporal pattern of resource supply did alter the intensity of competitive interactions among species. While additional studies in other systems are needed to evaluate the generality of these results, this study suggests that competitive intensity may not necessarily be a direct function of productivity or resource availability as traditionally assumed. Instead, the intensity of competitive interactions in resource-poor systems may depend upon the temporal pattern of resource supply.     

Differences in nitrate uptake of species from habitats rich or poor in nitrogen when grown at low nitrate concentrations,using a new growth technique

Summary A method is described for culturing plants at extremely low nutrient concentrations. Using a Braun infusion pump, a fixed amount of nitrate or ammonium was supplied continuously to plants growing in a culture vessel at a rate limiting the uptake of the plants. At a very low nitrogen concentration an equilibrium was established where uptake rate of the plants is equal to the rate of supply by the infusion pump. The nitrogen concentrations reached appeared to be in the order of 1 μM. The method compared the nitrate uptake byHypochaeris radicata L.ssp.radicata, H. radicata ssp.ericetorum Van Soest andUrtica dioica L. and ammonium uptake byH. radicata ssp.radicata andH. radicata ssp.ericetorum. Plants were cultivated in monocultures or in mixed cultures (two species per culture vessel). For the mixed cultures competition for nitrate (or ammonium) between the species was maintained for long periods. The capacities of the uptake systems of two subspecies ofH. radicata from places different in nitrogen supply and pH were adapted equally well to both low nitrate and low ammonium concentrations. Apparently factors other than nitrogen uptake play a part in the distribution of the subspecies. The capacity of the uptake system ofU. dioica, a nitrophilous species, was lower than that ofH. radicata ssp.radicata, a species from places poorer in nitrogen. This difference is related to the different distribution of the two species in the field. The present results are compared with those of previous experiments where K_m and V_max were measured and the significance of both parameters is discussed.     

Effects of Nitrogen and Salinity on Growth and Competition Between a Native Grass and an Invasive Congener

Numerous studies show that an increase in the availability of limiting resources can increase invasion by non-native plants into natural communities. One possible explanation is that the ability of natives to compete with non-natives tends to decrease when resource availability is increased. We tested this hypothesis in a competition experiment using two closely matched plant species and two environmental factors related to limiting resources in a coastal grassland system on Bodega Head in northern California. We grew the native grass Bromus carinatus and the non-native grass B. diandrus together and apart at different levels of soil nitrogen crossed with different levels of soil salinity. Both species are abundant in the grassland and previous work suggested that the abundance of B. carinatus is lower and the abundance of B. diandrus is higher on soil that has been enriched with nitrogen. Salinity has been shown to be negatively associated with invasion by B. diandrus into another California grassland, and to vary significantly over short distances in the grassland at Bodega Head, where it could affect water availability, which strongly limits plant growth during the dry season. Contrary to our prediction that low resource availabilities would increase the relative competitive ability of the native, the ability of B. carinatus to compete with B. diandrus was not greater when nitrogen availability was lower or when soil salinity was higher. Instead, high salinity increased the relative competitive ability of the non-native, and low nitrogen had little effect on competition. This suggests that preventing resource enrichment will not suffice to control invasion by non-native plant species in this grassland.     

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

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

Establishment of the invasive perennial herb Bunias orientalis L.: An experimental approach

A 2-year field experiment was performed to assess regeneration, establishment and growth of the invasive polycarpic perennial Brassicaceae Bunias orientalis L. from transplanted juveniles and propagules (seeds, fruits and root fragments). Further treatments were: different planting depths of the propagules, different root fragment sizes and a varied competition intensity by the matrix vegetation. In contrast to a high number of individuals established from planted juveniles (ca. 90 %), only relatively few propagules gave rise to seedlings or regenerating plants (0–16 %). However, upon emergence, all plants showed low mortality. Although the number of regenerated or recruited plants was affected by all treatments, only one treatment level (the use of very small root fragments of 1 cm length) completely failed in establishing plants. Plants experiencing low competition intensity attained high aboveground performance and reproductive output as well as reproductive success within two growth periods. Under high competition intensity recruitment, regeneration and growth were strongly negatively affected. The high morphological plasticity of rosette growth of B. orientalis seems to be advantageous for persistence in such situations. The results show that young B. orientalis plants are robust and can efficiently sequester and translate elevated resources into rapid growth and early reproductive output although the species is a rather long-lived iteroparous perennial. In addition, population foundation or regeneration can be based on both seeds and root fragments, even when buried under a thick soil layer. This seems to be a very efficient adaption to habitats which are subject to anthropogenic soil perturbations.     

向日葵种群中植株个体大小对其氮素利用策略的影响

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

Competition between the green alga Scenedesmus and the cyanobacterium Synechococcus under different modes of inorganic nitrogen supply

In this study, we evaluated growth responses of the green alga Scenedesmus and the cyanobacterium Synechococcus supplied with inorganic nitrogen in different ways. A competitive situation in which nitrogen was limiting was created in mixed cultures as well as in cultures growing in the same vessel but separated by a permeable dialysis membrane. Supplying inorganic nitrogen in small pulses at a high frequency favoured the cyanobacterium Synechococcus, whereas batch additions favoured the green alga Scenedesmus. When using a large-pulse/low-frequency supply mode, the yield of the green alga was higher when ammonium was added as nitrogen source compared to when nitrate was added. By contrast, the yield of the cyanobacterium was higher in the nitrate regime. However, uptake experiments using unialgal cultures showed that both organisms depleted the medium of ammonium more rapidly than they depleted the medium of nitrate; i.e. the higher yield of the cyanobacterium in the nitrate regime than in the ammonium regime can be attributed to the effects of competition with the green alga. Since nitrate assimilation involves the consumption of reductive power, we suggest that the outcome of competition was governed by the fact that green alga was light limited and therefore better able to compete for ammonium than for nitrate. The results from the laboratory studies are discussed in relation to results from an enclosure experiment performed in Lake Erken, Sweden. In that field experiment, in which additions of both phosphate and ammonium were applied every second day to 350-l enclosures, the green algal biomass increased exponentially during an incubation period of 22 days.     

Competition and resource availability in an annual plant community dominated by an invasive species,Carrichtera annua (L. Aschers.), in South Australia

The last decade has seen spirited debates about how resource availability affect the intensity of competition. This paper examines the effect that a dominant introduced species, Carrichtera annua, has upon the winter annual community in the arid chenopod shrublands of South Australia. Manipulative field experiments were conducted to assess plant community response to changing below-ground resource levels and to the manipulation of the density of C. annua. Changes in the density of C. annua had little effect on the abundance of all other species in the guild. Nutrient addition produced an increase in the biomass of the most abundant native species, Crassula colorata. An analysis of the root distribution of the main species suggested that the areas of soil resource capture of C. annua and C. colorata are largely segregated. Our results suggest that intraspecific competition may be stronger than interspecific competition, controlling the species responses to increased resource availability. The results are consistent with a two-phase resource dynamics systems, with pulses of high resource availability triggering growth, followed by pulses of stress. Smaller plants were nutrient limited under natural field conditions, suggesting that stress experienced during long interpulse phases may override competitive effects after short pulse phases. The observed differences in root system structure will determine when plants of a different species are experiencing a pulse or an interpulse phase. We suggest that the limitations to plant recruitment and growth are the product of a complex interplay between the length and intensity of the pulse of resource availability, the duration and severity of the interpulse periods, and biological characters of the species.     

Arbuscular mycorrhizal impacts on competitive interactions between Acacia etbaica and Boswellia papyrifera seedlings under drought stress

Aims Arbuscular mycorrhizal fungi can have a substantial effect on the water and nutrient uptake by plants and the competition between plants in harsh environments where resource availability comes in pulses. In this study we focus on interspecific competition between Acaia etbaica and Boswellia papyrifera that have distinctive resource acquisition strategies. We compared the extent of interspecific competition with that of intraspecific competition.Methods In a greenhouse study we examined the influence of Arbuscular Mycorrhiza (AM) and pulsed water availability on competitive interactions between seedlings of the rapidly growing species A. etbaica and the slowly growing species B. papyrifera. A factorial experimental design was used. The factors were AM, two water levels and five species combinationsImportant findings Seedlings of both species benefitted from AM when grown alone, and the positive growth response to pulsed water availability in B. papyrifera seedlings was in contrast with the negative growth response for A. etbaica seedlings. AM also affected the competitive performance of both species. B. papyrifera was not affected by intraspecific competition, whereas A. etbaica was negatively affected compared to the seedlings grown alone. This effect was stronger in the presence of AM. In interspecific competition, A. etbaica outcompeted B. papyrifera. Mycorrhiza and pulsed water availability did not affect the outcome of interspecific competition, and the aggressivity index of A. etbaica remained unchanged. The extent to which AM influences plant competition in a drought-stressed environment may depend on belowground functional traits of the species. AM and pulsed water availability could modify the balance between intraspecific and interspecific competition. By affecting the balance between intraspecific and interspecific competition, both factors could impact the establishment and survival of seedlings.     

Capture and allocation of nitrogen byQuercus douglasii seedlings in competition with annual and perennial grasses

Summary The spatial overlap of woody plant root systems and that of annual or perennial grasses promotes competition for soil-derived resources. In this study we examined competition for soil nitrogen between blue oak seedlings and either the annual grassBromus mollis or the perennial grassStipa pulchra under controlled outdoor conditions. Short-term nitrogen competition was quantified by injecting¹⁵N at 30 cm depth in a plane horizontal to oak seedling roots and that of their neighbors, and calculating¹⁵N uptake rates, pool sizes and¹⁵N allocation patterns 24 h after labelling. Simultaneously, integrative nitrogen competition was quantified by examining total nitrogen capture, total nitrogen pools and total nitrogen allocation.Stipa neighbors reduced inorganic soil nitrogen content to a greater extent than didBromus plants. Blue oak seedlings responded to lower soil nitrogen content by allocating lower amounts of nitrogen per unit of biomass producing higher root length densities and reducing the nitrogen content of root tissue. In addition, blue oak seedlings growing with the perennial grass exhibited greater rates of¹⁵N uptake, on a root mass basis, compensating for higher soil nitrogen competition inStipa neighborhoods. Our findings suggest that while oak seedlings have lower rates of nitrogen capture than herbaceous neighbors, oak seedlings exhibit significant changes in nitrogen allocation and nitrogen uptake rates which may offset the competitive effect annual or perennial grasses have on soil nitrogen content.     

Decrease in nitrate uptake and increase in proton release in zinc deficient cotton,sunflower and buckwheat plants

The effect of varied Zn supply on the pH of the nutrient solution and uptake of cations and anions was studied in cotton (Gossypium hirsutum L.), sunflower (Helianthus annuus L.) and buckwheat (Fagopyrum esculentum Moench) plants grown under controlled environmental conditions in nutrient solutions with nitrate as source of nitrogen. With the appearance of visual Zn deficiency symtoms, the pH of the nutrient solutions decreased from 6 to about 5 whereas the pH increased to about 7 when the plants were adequately supplied with Zn. In Zn deficient plants the pH decrease was associated with a shift in the cation-anion uptake ratio in favour of cation uptake. Of the major ions, uptake of Ca²⁺ and K⁺ was either not affected or only slightly lowered whereas NO₃ ^- uptake was drastically decreased in Zn deficient plants. Although the Zn nutritional status of plants hardly affected the NO₃ ^- concentrations in the plants, the leakage of NO₃ ^- from roots of Zn deficient plants into a diluted CaCl₂ solution was nearly 10 times higher than that of plants adequately supplied with Zn. In contrast to Zn deficiency, Mn deficiency in cotton plants neither affected NO₃ ^- uptake nor the pH of the nutrient solution.The results indicate that, probably as a consequence of the role of Zn in plasma membrane integrity and nitrogen metabolism, when Zn is deficient in dicotyledonous species net uptake of NO₃ ^- is particularly depressed which in turn results in an increase in cation-anion uptake ratio and a corresponding decrease in external pH. The ecological relevance of this rhizosphere acidification is discussed.     

Improving grasslands: the influence of soil moisture and nitrogen fertilization on the establishment of seedlings

1. In order to investigate the factors influencing the establishment of seedlings in permanent grassland, the influence of soil moisture and nitrogen fertilization on competition between established plants of Lolium perenne and seedlings of Phleum pratense or Trifolium pratense was studied in two experiments under greenhouse conditions using the 'split-box'-technique.
2. There was no difference in the production of plant dry matter of P. pratense or T. pratense between 30% volumetric soil water content (−0·005 MPa) and 22% (−0·04 MPa), but 15% soil moisture (−0·33 MPa) reduced plant growth. L. perenne yields were linearly reduced by reduced soil moisture content.
3. Shoot competition from L. perenne reduced the plant dry matter yield of P. pratense and T. pratense more than did root competition in these experiments. When shoot competition was present, differences between moisture contents were not detected, indicating that light was probably the limiting resource under such conditions. No significant interaction between root competition and soil moisture was observed for plant weight.
4. Root competition was not prevented even though sufficient water and nitrogen were supplied. This indicated either that some other growth factor was limiting or the plants competed for resources at the root hair level even though sufficient resources were supplied at the pot or field scale. Therefore, in the situation of direct drilling of species during grassland renovation, it may be difficult to alleviate competition by adequate provision of water and nitrogen.     

Resource pulses of dead periodical cicadas increase the growth of American bellflower rosettes under competitive and non-competitive conditions

This experiment investigates how pulsed nutrient resources interact with intraspecific competition to influence biomass production and nutrient use efficiency in the American bellflower (Campanulastrum americanum). The competitive environment of these plants was manipulated by growing plants alone or with neighboring conspecifics, and the occurrence of pulsed nutrient resources was manipulated through the addition of 17-year periodical cicada (Magicicada septendecim) carcasses in order to simulate naturally occurring pulsed resources in this system. The addition of cicada carcasses increased mean plant biomass by 61 % compared to non-supplemented plants, while competition decreased mean plant biomass by 44 % compared to plants grown without competition; these effects were additive. In comparison, nutrient use efficiency decreased in fertilized plants (cicada-supplemented plants showed 20 % greater foliage nitrogen concentrations compared to non-supplemented plants), but was not significantly affected by the plant’s competitive environment. In addition, cicada supplementation did not significantly increase the biomass asymmetry in competing pairs of plants. These results suggest that these plants increased their total nutrient uptake at a timescale commensurate with the pulsed increase in nutrient availability due to cicada carcass decomposition.     

Competition between a toxic and a non-toxic <Emphasis Type="Italic">Microcystis</Emphasis> strain under constant and pulsed nitrogen and phosphorus supply

The toxicity of a harmful algal bloom is strongly determined by the relative abundance of non-toxic and toxic genotypes and might therefore be regulated by competition for growth-limiting resources. Here, we studied how the toxic Microcystis aeruginosa strain PCC 7806 and a non-toxic mutant compete for nitrogen and phosphorus under constant and pulsed nutrient supply. Our monoculture and competition experiments show that these closely related genotypes have distinct nutrient physiologies and that they differ in their ability to compete for nitrogen and phosphorus. The toxic wild type won the competition under nitrogen limitation, while the non-toxic mutant dominated under phosphorus limitation. Pulses of both nitrogen and phosphorus increased the dominance of the toxic genotype, which lead to an even faster competitive exclusion of the non-toxic genotype under nitrogen pulses and to coexistence of both genotypes under phosphorus pulses. Our findings indicate that the genotype level dynamics driven by resource competition can be an important factor in determining cyanobacterial bloom toxicity.     

Uptake and transport of N,P and K in tomato supplied with nettle water and nutrient solution

Water extract of stinging nettle (Urtica dioica) has a growth stimulating effect on plants. This investigation elucidated effects of nettle water on uptake and transport of N, P and K. Tomato plants (Solanum lycopersicum L. cv. Dansk export) were grown in sand culture 6–8 weeks. Plants were supplied with nettle water and nutrient solution was used as a control medium. Uptake and transport of N, P and K⁺ were determined with isotopes (¹⁵N,³²P and⁸⁶Rb⁺ as a tracer for K⁺) and ion-selective electrodes and in exudation experiments. A 15% higher uptake of nitrogen (¹⁵N assay) was found after nettle water treatment compared with the nutrient solution control. The total amount of nitrogen was also higher in plants cultivated with nettle water. Transport of inorganic and organic nitrogen, measured in exudation experiments, was more than 50% higher for plants supplied with nettle water compared with plants supplied with nutrient solution. In contrast, nettle water had no effect on uptake, transport or total amount of phosphorus and potassium in the plants. Experiments in hydroculture showed that nettle water had a strong pH-elevating effect. Uptake of NH ₄ ⁺ was strongly stimulated by nettle water compared with nutrient solution. By holding pH at a constant level during the uptake period for 6 h, the uptake of NH ₄ ⁺ from nettle water was significantly lower when no adjustment of pH was made. Consequently a good deal of the NH ₄ ⁺ uptake enhancement by nettle water could be explained by pH-stimulation. Assays with the uncoupler/inhibitor 2,4-dinitrophenol (DNP) and dichlorophenyl-dimethyl-urea (DCMU) showed that uptake of nitrogen from nettle water was less metabolically-linked than uptake from a corresponding nutrient solution. All together, nettle water seems to stimulate the uptake of nitrogen, but not phosphorus or potassium.     

Interspecific competition enhances nitrogen fixation in an actinorhizal shrub

In forest understory restoration, the establishment of reintroduced species may be strongly linked to their ability to compete for belowground resources. In this study, we provide isotopic and morphological evidence for competition-induced increases in nitrogen fixation by Morella cerifera (L.) Small (wax myrtle) when planted with Pinus palustris Mill (longleaf pine). Compared to a competition-free treatment, we found no significant differences in tissue N concentrations for M. cerifera. However, ¹⁵N enrichment in leaves, stems and roots, as well as whole-plant values for nitrogen derived from fertilizer were significantly lower when the plants were subject to interspecific competition from P. palustris. Plants in the competition treatment also allocated a significantly greater percentage of belowground biomass to root nodules than those in the competition-free treatment (0.65 vs. 0.41%). This strongly suggests that M. cerifera is capable of upregulating nitrogen fixation in response to interspecific competition. This may help explain why M. cerifera outperformed non-nitrogen-fixing species reintroduced on the same site.     

Effects of collembola (arthropoda) and relative germination date on competition between mycorrhizalPanicum virgatum (Poaceae) and non-mycorrhizalBrassica nigra (Brassicaceae)

In this experiment the separate and interactive effects of grazing of vesicular-arbuscular mycorrhizal hyphae by collembola and of relative germination date on competition between the mycorrhizal perennial grassPanicum virgatum and the non-mycorrhizal annual cruciferBrassica nigra were investigated. In the absence of competition,P. virgatum mass and P uptake were not affected by collembola grazing; grazing did reduce tissue N concentration and root: shoot ratio. Competition fromB. nigra plants of the same age/size (“simultaneous competition”) significantly reducedP. virgatum total, root, and shoot mass relative to control plants not subject to competition. In contrast, when in competition,B. nigra plants did not differ in biomass fromB. nigra controls grown without competition. Simultaneous competition also reduced N and P uptake byP. virgatum, but not byB. nigra. Grazing by collembola during simultaneous competition increased the differences in nutrient uptake and tended to shift the competitive balance further toward the non-mycorrhizalB. nigra. WhenP. virgatum plants were subjected to competition fromB. nigra plants which germinated three weeks later (“offset competition”) the situation was reversed: offsetB. nigra plants were negatively affected by competition while the larger, olderP. virgatum plants were not. Thus, relative germination date is important in determining the relative competitive ability of these two species. Grazing by collembola did not affect offset competition. The grazing of VAM hyphae by collembola appears to increase N availability in this experimental system. Under simultaneous competition, this N is taken up by the more extensive root system ofB. nigra; under offset conditions, the root system of the smallerB. nigra plants is insufficient to take advantage of the added resources. Thus, we suggest that grazing-induced transient changes in nutrient availability and the differential abilities of the two species to make use of these added resources constitute the mechanisms by which relative germination date and collembola grazing influence competition.     

Shading and the capture of localized soil nutrients: nutrient contents,carbohydrates, and root uptake kinetics of a perennial tussock grass

Summary The ability to exploit spatial and temporal heterogeneity in soil resources can be one factor important to the competitive balance of plants. Competition above-ground may limit selective plant responses to below-ground heterogeneity, since mechanisms such as root proliferation and alterations in uptake kinetics are energy-dependent processes. We studied the effect of shading on the ability of the perennial tussock grassAgropyron desertorum to take up nutrients from enriched soil microsites in two consecutive growing seasons. Roots of unshaded plants selectively increased phosphate uptake capacity in enriched soil microsites (mean increases of up to 73%), but shading eliminated this response. There were no changes in ammonium uptake capacity for roots in control and enriched patches for either shaded or unshaded plants. The 9-day shade treatments significantly reduced total nonstructural carbohydrate (TNC) concentrations for roots in 1990, but had no apparent effect on root carbohydrates in 1991 despite dramatic reductions in shoot TNC and fructan concentrations. Enrichment of the soil patches resulted in significantly greater phosphate concentrations in roots of both shaded and unshaded plants, with less dramatic differences for nitrogen and no changes in potassium concentrations. In many respects the shaded plants did surprisingly well, at least in terms of apparent nutrient acquisition. The effects of aboveground competition on nutrient demand, energy requirements, and belowground processes are discussed for plants exploiting soil resource heterogeneity.     

A Mechanistic Study of Plant and Microbial Controls over R* for Nitrogen in an Annual Grassland

Differences in species'' abilities to capture resources can drive competitive hierarchies, successional dynamics, community diversity, and invasions. To investigate mechanisms of resource competition within a nitrogen (N) limited California grassland community, we established a manipulative experiment using an R* framework. R* theory holds that better competitors within a N limited community should better depress available N in monoculture plots and obtain higher abundance in mixture plots. We asked whether (1) plant uptake or (2) plant species influences on microbial dynamics were the primary drivers of available soil N levels in this system where N structures plant communities. To disentangle the relative roles of plant uptake and microbially-mediated processes in resource competition, we quantified soil N dynamics as well as N pools in plant and microbial biomass in monoculture plots of 11 native or exotic annual grassland plants over one growing season. We found a negative correlation between plant N content and soil dissolved inorganic nitrogen (DIN, our measure of R*), suggesting that plant uptake drives R*. In contrast, we found no relationship between microbial biomass N or potential net N mineralization and DIN. We conclude that while plant-microbial interactions may have altered the overall quantity of N that plants take up, the relationship between species'' abundance and available N in monoculture was largely driven by plant N uptake in this first year of growth.     

INTERACTIONS BETWEEN PULSED NUTRIENT SUPPLIES AND A PHOTOCYCLE AFFECT PHYTOPLANKTON COMPETITION FOR LIMITING NUTRIENTS IN LONG-TERM CULTURE1

The influence of periodic nutrient supplies and a photocycle on phytoplankton competition for limiting nutrients was examined using the diatoms Thalassiosira rotula Meunier (clone 411) and Chaetoceros sp. cf. vixvisibilis Schiller (clone 847). Chaetoceros sp. cf. vixvisibilis displaced T. rotula from ammonium-limited cultures under constant light irrespective of whether ammonium was supplied continuously, in 6 pulses.day^?1 or in a single daily pulse. In contrast, the species coexisted under the 14:10 h LD photocycle under either continuous or pulsed ammonium supplies with the relative abundance of C. sp. cf. vixvisibilis increasing as the interval between ammonium additions lengthened. Coexistence was not observed with either silicic acid or nitrate limitation. Chaetoceros sp. cf. vixvisibilis displaced T. rotula from both nitrate- and silicic acid-limited chemostat cultures and from semi-continuous cultures grown under the same photoperiod that produced coexistence with a daily pulse of ammonium. The presence of a photocycle was both necessary and sufficient to permit coexistence with ammonium limitation. Under continuous ammonium supply the photocycle may have induced a temporal separation of ammonium uptake between species, permitting sharing of the limiting nutrient and coexistence. In contrast, the species were shown to be in direct competition for the daily ammonium pulse. A competition model suggested that coexistence in this case arose from a balance between the species’ammonium uptake rates and their nitrogen demands for steady-state growth induced by the photocycle. The results indicate that variations in nutrient supply rates may contribute to the coexistence of phytoplankton species in the sea, but that the identity of the limiting nutrient and the influence of variations in other non-limiting resources play important roles in affecting the outcome of nutrient competition among planktonic algae.