EVOLUTION OF BRASSICA RAPA L. (CRUCIFERAE) POPULATIONS IN INTRA- AND INTERSPECIFIC COMPETITION

Populations of Brassica rapa were grown for three generations in each of two environments: intraspecific competition, with four surrounding Brassica rapa neighbors per pot, and interspecific competition, with two Raphanus sativus neighbors per pot. In each environment, the largest (by flower number) 10% of the plants were outcrossed and provided seeds for the next generation. As a control, a randomly chosen 10% of the plants in each environment were outcrossed to produce seed for the next generation. Each of these four treatments, the selected lines in intra- and interspecific competition and the corresponding control lines, was maintained for three generations. After a single generation of growth in a common, no-competition environment, replicate plants from each treatment were grown with no competition and with intra- and interspecific competition for determination of growth responses. After two generations of selection, flower number in the intraspecific-selection line had increased by more than 50% over that in the control line and by more than 19% over that under interspecific selection. After a common-environment generation, plants from the intraspecific-selection line were shown to have significantly faster growth in height and flower number as seedlings. Plants in the interspecific-selection line showed similar but nonsignificant trends. No differences in seed mass, emergence time, or photosynthetic rate were found between control and selected lines in either intra- or interspecific competition. Some differences between control and selected lines were noted in biomass allocation related to differences in phenology. The results demonstrate that performance in competitive environments can evolve through changes in plant development but that rates of evolution will differ in intra- and interspecific competition.     

Competition and genetic background in a rapid-cycling cultivar of brassica rapa (Brassicaceae)

We examined the effects of a slight genetic disadvantage in a competitive situation by comparing the performance of inbred and outcrossed Brassica rapa plants over a range of neighbor densities, using a rapid-cycling, self-compatible cultivar of this species. We also examined the genetic correlations in performance among plants grown alone and with intra- and interspecific competitors. Competition had a strong effect on biomass and on the number of flowers produced, but differences in biomass between inbred and outcrossed plants were dependent on the competitive environment. Outcrossed plants outperformed those that were selfed only at intermediate densities of neighbors; at high densities and in the absence of competition inbred and outcrossed plants did not differ. For outcrossed families, performance without competitors and in intra- and interspecific competition were all highly correlated, but for selfed families, correlations were low, and performance in competition was not predictable from that of plants grown alone. Thus, the phenotypic expression of genetic differences may depend on the density of neighbors with which plants are grown.     

Differential effect of inter- and intraspecific competition on the performance of invasive and native Taraxacum species

Inter- and intraspecific competitive abilities are significant determinants of invasive success and the ecological impact of non-native plants. We tested two major hypotheses on the competitive ability of invasive species using invasive (Taraxacum officinale) and native (T. platycarpum) dandelions: differential interspecific competitive ability between invasive and native species and the kin recognition of invasive species. We collected seeds from two field sites where the two dandelion species occurred nearby. Plants were grown alone, with kin (plants from the same maternal genotype) or strangers (plants from different populations) of the same species, or with different species in a growth chamber, and the performance at the early developmental stage between species and treatments was compared. The invasive dandelions outcompeted the native dandelions when competing against each other, although no difference between species was detected without competition or with intraspecific competition. Populations of native species responded to interspecific competition differently. The effect of kinship on plant performance differed between the tested populations in both species. A population produced more biomass than the other populations when grown with a stranger, and this trend was manifested more in native species. Our results support the hypothesis that invasive plants have better competitive ability than native plants, which potentially contributes to the establishment and the range expansion of T. officinale in the introduced range. Although kin recognition is expected to evolve in invasive species, the competitive ability of populations rather than kinship seems to affect plant growth of invasive T. officinale under intraspecific competition.     

Effect of arbuscular mycorrhiza on inter- and intraspecific competition of two grassland species

We were interested in the role of arbuscular mycorrhiza (AM) in the competition between plants of different sizes. A pot experiment of factorial design was established, in which AM root colonization and competition were used as treatments. Five-week-old Prunella vulgaris seedlings were chosen as target plants (i.e. plants whose response to competition was studied) and the following (13 replicates of each) were used as neighbours: (1) a large, 10-week-old P. vulgaris, (2) two P. vulgaris seedlings, and (3) a large, 10-week-old Fragaria vesca. In the experiment where small neighbours were grown together with small target plants, competition did not reduce target plant weight significantly, compared to the other two treatments. The competitive effects of large neighbours were significant, regardless of species (both older neighbours reduced the weights of target plants similarly), but there was a clear difference between intra- and interspecific competition when plants were mycorrhizal. In intraspecific competition with a large neighbour, the target plant shoot weight was reduced 24% when inoculated with AM. Thus, AM amplified rather than balanced intraspecific competition. In interspecific competition with old F. vesca, the shoot weights of target plants were 22% greater when inoculated with AM than when non-mycorrhizal. The results showed that, for given soil condition, AM might increase species diversity by increasing competitive intraspecific suppression and decreasing the interspecific suppression of small plants by larger neighbours.     

Integrated response of intercropped maize and potatoes to heterogeneous nutrients and crop neighbours

Background and Aims

In communities, plants often simultaneously interact with intra- and inter-specific neighbours and heterogeneous nutrients. How plants respond under these conditions and then affect the structure and function of communities remain important questions.

Methods

Maize (Zea mays L.) was intercropped with potatoes (Solanum tuberosum L.). In the field experiment, we applied fertilizer both homogeneously and heterogeneously under monocropping and intercropping conditions. The heterogeneous nutrient treatment in intercropping was designed with different fertilizer placements, at intraspecific and interspecific rows, respectively. In the pot experiment, crops were grown under both homogeneous and heterogeneous nitrogen conditions with single plant, intraspecific and interspecific competition. Shoot and root biomass and yield were measured to analyse crop performance.

Results

In the field experiment, the heterogeneous nitrogen, compared with the homogenous one, enhanced the performance of the intercropped crop. Importantly, this effect of heterogeneous nitrogen was greater when fertilizer was applied at interspecific rows, rather than at intraspecific rows. Moreover, in pot experiments, the root foraging precision of the two crops was increased by interspecific neighbours, but only that of potatoes was increased by intraspecific neighbours.

Conclusions

The integrated responses of plants to heterogeneous neighbours and nutrients depend on the position of nutrient-rich patches, which deepen our understanding of the function of plant diversity, and show that fertilizer placement within multi-cropping systems merits more attention. Moreover, the enhanced utilization of heterogeneous nitrogen could drive overyielding in multi-cropping systems.     

INTRAPOPULATION DIFFERENTIATION IN ANNUAL PLANTS. III. THE CONTRASTING EFFECTS OF INTRA- AND INTERSPECIFIC COMPETITION

The roles of intraspecific and interspecific competition in producing differentiation within populations of Veronica peregrina were studied in two populations under controlled, greenhouse conditions. In nature, each population spans an environmental gradient across the center and sides of a temporary, vernal pool in California. Individuals at the center are subjected to intense intraspecific competition produced by high densities (to 30 seedlings/cm²) generated by quasi-simultaneous germination (90% of seeds germinate in one week). Individuals at the periphery are subjected to interspecific competition with grasses, which shade out the Veronica 4–6 weeks after the onset of winter growth. I predicted that 1) when grown under immediate intraspecific competition in the greenhouse, offspring of plants from the central subpopulation (C) would perform better (i.e., grow larger and produce more seeds) than those from the periphery (P) and that 2) when grown under delayed interspecific competition provided by Agrostis tenuis and Lollium multiflorum, offspring of plants from the periphery would perform better than those from the center. Both predictions were confirmed. The center-periphery differences were pronounced and statistically significant in an undisturbed population (V-2), while in a population disturbed by yearly plowing (V-3), the differences tended to be consistent with those in V-2 but seldom significant. Distribution of variability tended to be positively skewed and/or leptokurtic in subpopulations grown under “foreign” competition (i.e., intraspecific for P plants and interspecific for C plants) but was normally distributed following exposure to “familiar” competition. Timing of competition affected many results. There were four additional significant differences between the central and peripheral subpopulations. 1) Germination rate: the faster rate in central plants can be advantageous under immediate intraspecific competition. The slower rate in peripheral plants can be advantageous under conditions of erratic and unpredictable soil moisture. 2) Response to nutrient competition: central plants were more sensitive to N-deficiency and peripheral plants were more sensitive to P-deficiency. 3) Allocation of biomass: central plants allocated a greater proportion of biomass to seeds, while peripheral plants allocated a greater proportion of biomass to leaves under all growing conditions. 4) Root elongation: at the seedling stage, central plants have longer roots, while at the adult stage, peripheral plants have longer roots (but not more root mass). Most components of this complex pattern of differentiation are interpretable in an adaptive context. Other results defy simple explanations and underline the importance of phenotypic plasticity, which was pronounced in the competition experiments.     

Interspecific and intraspecific interactions between salt marsh plants: integrating the effects of environmental factors and density on plant performance

There has been much debate about the role of plant interactions in the structure and function of vegetation communities. Here the results of a pot experiment with controlled environments are described where three environmental variables (nutrients, sediment type and waterlogging) were manipulated factorially to identify their effects on the growth and intensity of interactions occurring between Spartina anglica and Puccinellia maritima. The two species were grown in split-plot planting treatments, representing intraspecific and interspecific addition series experiments, to determine individual and interactive effects of environmental factors and plant interactions on plant biomass.
Above-ground growth of both species involved interactions between the environmental and planting treatments, while below-ground, environmental factors affected the biomass irrespective of planting treatments. It was suggested that this difference in growth response is evidence that in our experiment plant interactions between the two species occur primarily at the above-ground level.
The intensity of plant interactions varied in a number of ways. First, interactions between Spartina and Puccinellia were distinctly asymmetrical, Puccinellia exerting a competitive effect on Spartina, with no reciprocal effect, and with a facilitative effect of Spartina on Puccinellia in low nutrient conditions. Second, the interactions varied in intensity in different environmental conditions. Interspecific competitive effects of Puccinellia on Spartina were more intense in conditions favourable to growth of Puccinellia and reduced or non-existent in environments with more abiotic stress. Third, intraspecific competition was found to be less intense for both species than interspecific interactions. Finally, the intensity of plant interactions involving both species was more intense above ground than below ground, with a disproportionate reduction in the intensity of interspecific competition below relative to above ground in treatments with less productive sediments and greater immersion. This is interpreted as reflecting a potential mechanism by which Spartina may be able to evade competitive neighbours.     

Competitive interactions are mediated in a sex‐specific manner by arbuscular mycorrhiza in Antennaria dioica

• Plants usually interact with other plants, and the outcome of such interaction ranges from facilitation to competition depending on the identity of the plants, including their sexual expression. Arbuscular mycorrhizal (AM) fungi have been shown to modify competitive interactions in plants. However, few studies have evaluated how AM fungi influence plant intraspecific and interspecific interactions in dioecious species.
• The competitive abilities of female and male plants of Antennaria dioica were examined in a greenhouse experiment. Females and males were grown in the following competitive settings: (i) without competition, (ii) with intrasexual competition, (iii) with intersexual competition, and (iv) with interspecific competition by Hieracium pilosella – a plant with similar characteristics to A. dioica. Half of the pots were grown with Claroideoglomus claroideum, an AM fungus isolated from the same habitat as the plant material. We evaluated plant survival, growth, flowering phenology, and production of AM fungal structures.
• Plant survival was unaffected by competition or AM fungi. Competition and the presence of AM fungi reduced plant biomass. However, the sexes responded differently to the interaction between fungal and competition treatments. Both intra‐ and interspecific competition results were sex‐specific, and in general, female performance was reduced by AM colonization. Plant competition or sex did not affect the intraradical structures, extraradical hyphae, or spore production of the AM fungus.
• These findings suggest that plant sexual differences affect fundamental processes such as competitive ability and symbiotic relationships with AM fungi.

     

喜旱莲子草和接骨草竞争对模拟增温的响应北大核心CSCD

研究外来入侵植物与本地植物种竞争对气候变暖的响应,对于预测未来气候变化背景下入侵植物的入侵趋势、理解其入侵机制以及筛选生态替代种具有重要的意义。以入侵我国的外来植物喜旱莲子草(Alternanthera philoxeroides)和本地植物种接骨草(Sambucus chinensis)为材料,通过两种植物单栽、纯栽和混栽,采用红外辐射加热器模拟增温,研究了两种植物竞争对模拟增温的响应。结果表明:(1)在模拟增温期间(2013年5–12月),增温组空气平均温度比不增温组提高了0.47℃,相对湿度降低了1.87%;(2)混栽的喜旱莲子草除根冠比与单栽无显著差异外,其余各生物量和根系形态指标均显著低于单栽喜旱莲子草;无竞争、种间竞争和种内竞争三种竞争间,接骨草除根冠比、细根与总根生物量比、比根长和比根表面积无显著差异外,其余指标均呈现无竞争>种间竞争>种内竞争的趋势;(3)无竞争、种间竞争和种内竞争三种条件下,喜旱莲子草各指标在增温和不增温处理间差异均不显著,而接骨草总生物量和根生物量在无竞争和种间竞争条件下增温处理均显著低于不增温处理,在种内竞争条件下则相反;(4)增温使接骨草的相对拥挤系数降低,接骨草对温度升高反应敏感,而喜旱莲子草则表现出一定的适应性。由此推测,在中度遮阴陆生生境中,接骨草有望成为喜旱莲子草生物替代控制的材料。     

Competition and facilitation within and between a snail and a mayfly larva and the effect on the grazing process

We studied the competitive effects within and between two taxonomically distant freshwater herbivores, a snail and a mayfly, common in Swedish lakes, Lymnaea peregra and Cloeon dipterum, respectively, and their effect on grazing in a laboratory experiment. The experimental set-up consisted of 2-l aquaria, each containing a periphyton covered tile. Intra- and interspecific effects were tested by increasing the density of one species at a time in four different treatments, (1) snails (intraspecific treatment), (2) mayflies (intraspecific treatment), (3) mixed-snails (interspecific treatments, snails kept constant) and (4) mixed-mayflies (interspecific treatments, mayflies kept constant). Intraspecific competition affected both snails and mayflies negatively, i.e. increasing mortality with increasing con-specific density. Furthermore, there was a decrease in snail growth with increasing snail density. In the mixed-species treatments both species changed their microhabitat use indicating interspecific competition. Despite this, we also found a positive effect of mayfly density on snail growth, most likely due to indirect commensalism. No density-dependent effect of grazing on periphyton was found, probably due to interference competition between grazers. However, there was a significant difference in periphyton biomass, due to species composition of grazers. Irrespective of their densities, if they co-existed, the two grazer species decreased the periphyton biomass significantly compared with both single-species treatments. We considered this as a joint action of facilitation and interaction. Our results suggest that competition can be an important structuring factor in macroinvertebrate communities and that species composition can be significant for ecosystem processes within lentic environments.     

Spatial aggregation facilitates coexistence and diversity of wild plant species in field margins

European agri-environment schemes encourage farmers to establish sown field margin strips to protect and enhance wild plant diversity. However, plant diversity in such wild plant sowings based on seed mixtures is often low due to the high competitiveness of few, common species. Here we analysed whether intraspecific aggregation could enhance the performance of less competitive species, and how plant performance is influenced by the number of species in a mixture. We focused on inter- and intraspecific competition between six agricultural wild plant species (Centaurea cyanus, Calendula arvensis, Melilotus officinalis, Poa annua, Bromus mollis, Medicago lupulina), and tested (i) two different seeding patterns (intraspecifically aggregated vs. randomly dispersed) and (ii) three different species mixtures (monocultures, three-species, and six-species mixtures). Plant performance was measured in terms of number of individuals, biomass per individual, and biomass per m². Intraspecific aggregation resulted in higher numbers of individuals of all species, while mixtures generated lower numbers of individuals. The performance of plant species differed depending on their position in the competitive hierarchy. Competitively weak species suffered much less from intraspecific than interspecific competition in terms of biomass, and the competitively weakest species became even excluded in the most species rich and randomly dispersed sowings with high interspecific competition. In conclusion, the performance of wild plant species was influenced by both seeding pattern and number of species in a mixture. Intraspecific aggregation enabled the coexistence of competitively weak species by reducing interspecific competitive exclusion processes. Consequently, agri-environmental schemes designed to preserve and enhance biodiversity should consider small-scale processes influencing the distribution and abundance of plants, and develop new agricultural sowing technologies to cultivate competitively weak and endangered wild plant species.     

Pollination of Ipomopsis aggregata (Polemoniaceae): effects of intra- vs. interspecific competition

Although plants may simultaneously experience intra- and interspecific competition for pollination, their relative strength has rarely been experimentally evaluated. Yet because intra- and interspecific competition can be caused by different mechanisms, their effect on the ecology and evolution of plants may differ. To determine the relative strength of intra- and interspecific competition for pollination, I manipulated the presence of heterospecifics and density of conspecifics using Ipomopsis aggregata as the focal species. All plots contained I. aggregata and Castilleja linariaefolia, but C. linariaefolia inflorescences were removed from half of the plots to create the heterospecifics-absent treatment. Within each plot, all I. aggregata inflorescences were removed from a 5-m radius around a focal plant to create a low conspecific density experimental unit, and a group of 12 I. aggregata plants/1 m² was designated as a high conspecific density unit. Conspecific pollen deposition was reduced when C. linariaefolia was present but was not influenced by I. aggregata density. Although seed set per fruit was reduced by 17% when C. linariaefolia was present, it was not significantly influenced by either treatment. Interspecific competition for pollination is stronger than intraspecific competition in the I. aggregata–C. linariaefolia system, but neither process appears to influence plant fitness.     

Variation in mycorrhizal growth response influences competitive interactions and mechanisms of plant species coexistence

Plant species vary in their growth response to arbuscular mycorrhizal (AM) fungi, with responses ranging from negative to positive. Differences in response to AM fungi may affect competition between plant species, influencing their ability to coexist. We hypothesized that positively responding species, whose growth is stimulated by AM fungi, will experience stronger intraspecific competition and weaker interspecific competition in soil containing AM fungi, while neutrally or negatively responding species should experience weaker intraspecific and stronger interspecific competition. We grew Plantago lanceolata, which responds positively to AM fungi, and Bromus inermis, which responds negatively to AM fungi, in an additive response surface competition experiment that varied the total density and relative frequency of each species. Plants were grown in sterilized background soil that had been inoculated with whole soil biota, which includes AM fungi, or a microbial wash, that contained other soil microbes but no AM fungi, or in sterilized soil that contained no biota. The positively responding P. lanceolata was more strongly limited by intraspecific than interspecific competition when AM fungi were present. By contrast, the presence of AM fungi decreased the strength of intraspecific competition experienced by the negatively responding B. inermis. Because AM fungi are almost always present in soil, strong intraspecific competition in positively responding species would prevent them from outcompeting species that respond neutrally or negatively to AM fungi. The potential for increased intraspecific competition to offset growth benefits of AM fungi could, therefore, be a stabilizing mechanism that promotes coexistence among plant species.     

意大利苍耳与乌拉尔甘草种间竞争能力比较

意大利苍耳已经开始入侵乌拉尔甘草农田,然而对其入侵后果目前知之甚少。通过研究意大利苍耳与乌拉尔甘草的种间竞争关系,以期为意大利苍耳对乌拉尔甘草农田生态系统的入侵能力及入侵后果的评估提供试验依据。模拟了甘草农田的土壤水肥条件,采用取代试验法,设置意大利苍耳与乌拉尔甘草的单种种植和混种种植2种种植模式,待意大利苍耳生育期结束后进行收获,分别比较了该两种植物的个体生长及生物量积累在单种种植与混种种植两种处理间的差异,并比较单种和混种模式下乌拉尔甘草的地下器官中甘草酸含量的差别,通过计算相对产量、相对竞争强度和竞争攻击系数来比较该两种植物种间竞争能力的相对强弱。结果表明：与单种模式相比,混种模式下意大利苍耳个体生长的形态学性状和有性繁殖能力均有显著增加,其株高、冠幅和种子数量较单种处理分别增加了13%、27%和56%。而乌拉尔甘草的个体生长及克隆繁殖能力均显著降低,其根瘤的形成也受到显著的抑制,混种处理的乌拉尔甘草的株高、冠幅、根总长度、根总表面积、根平均直径以及根瘤数量比单种处理分别下降了35%、45%、55%、63%、19%和76%。单种处理下每株甘草根状茎的平均条数为3条,而与意大利苍耳混种后,其根状茎的发育被完全抑制。与单种处理相比,混种处理中的意大利苍耳生物量积累均显著的增加了,其中根、茎、叶、果实及总生物量与单种模式相比分别增加了84%、73%、84%、73%和77%,而混种模式却极显著降低了乌拉尔甘草生物量和地下器官甘草酸含量的积累,使其根、茎、叶、总生物量以及甘草酸含量与单种模式相比分别下降了72%、80%、65%、71%和63%。混种模式中的意大利苍耳相对产量（RY_a）大于1,而乌拉尔甘草相对产量（RY_b）小于1,表明意大利苍耳受到邻株同种其他个体的种内竞争压力大于来自邻株乌拉尔甘草的种间竞争压力,而乌拉尔甘草受到邻株同种其他个体的种内竞争压力则小于来自邻株意大利苍耳的种间竞争压力。混种模式中的意大利苍耳相对竞争强度（RCI_a）小于0,其竞争攻击系数（A_q）大于0,而乌拉尔甘草的相对竞争强度（RCI_b）则在0-1之间,其竞争攻击系数（A_b）小于0,表明在该两种植物的混生群落中,乌拉尔甘草的竞争能力弱于意大利苍耳。总体来看,在二者混生的群落中,意大利苍耳在竞争中占据明显的优势地位,对乌拉尔甘草的产量和品质均造成强烈的负面影响。     

Interspecific competition changes reproductive output but does not increase reproductive costs in a grassland perennial

In plants, it is hypothesized that allocation trade-offs may appear only when expenditures like seed production are high or external resources are scarce. In this study, we tested whether reproductive costs are more pronounced under enhanced interspecific competition.In a common garden, we investigated phenotypic correlations between sexual reproduction, clonal growth and storage structures in the grassland perennial, Succisa pratensis. During the past 50 years, habitats of this species have faced an expansion of clonal grasses that increase competition intensity. We simulated this process by growing five populations of Succisa from high- and low-production habitats with its clipped and non-clipped competitor, Agrostis capillaris. In addition, we experimentally removed flower heads of Succisa plants from one population grown with and without A. capillaris.We demonstrated costs of sexual reproduction by flower-head removal (resulting in increased plant size and relative allocation to belowground structures) but not by phenotypic correlations. We found no evidence that reproductive costs increase in a competitive environment and the opposite pattern was shown in both approaches used. However, high competition intensity reduced relative investment to flower-head production. In plants from low-production habitats, competition also reduced the absolute number of flower heads and belowground biomass as a result of smaller plant size. We assume that populations from low-production habitats are more prone to extinction as they have a reduced likelihood of local persistence and of escape to more suitable habitats during advancing succession.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Competition increasingly dominates the responsiveness of juvenile beech and spruce to elevated CO2 and/or O3 concentrations throughout two subsequent growing seasons

Saplings of Fagus sylvatica and Picea abies were grown in mono‐ and mixed cultures in a 2‐year phytotron study under all four combinations of ambient and elevated ozone (O₃) and carbon dioxide (CO₂) concentrations. The hypotheses tested were (1) that the competitiveness of beech rather than spruce is negatively affected by the exposure to enhanced O₃ concentrations, (2) spruce benefits from the increase of resource availability (elevated CO₂) in the mixed culture and (3) that the responsiveness of plants to CO₂ and O₃ depends on the type of competition (i.e. intra vs. interspecific). Beech displayed a competitive disadvantage when growing in mixture with spruce: after two growing seasons under interspecific competition, beech showed significant reductions in leaf gas exchange, biomass development and crown volume as compared with beech plants growing in monoculture. In competition with spruce, beech appeared to be nitrogen (N)‐limited, whereas spruce tended to benefit in terms of its plant N status. The responsiveness of the juvenile trees to the atmospheric treatments differed between species and was dominated by the type of competition: spruce growth benefited from elevated CO₂ concentrations, while beech growth suffered from the enhanced O₃ regime. In general, interspecific competition enhanced these atmospheric treatment effects, supporting our hypotheses. Significant differences in root : shoot biomass ratio between the type of competition under both elevated O₃ and CO₂ were not caused by readjustments of biomass partitioning, but were dependent on tree size. Our study stresses that competition is an important factor driving plant development, and suggests that the knowledge about responses of plants to elevated CO₂ and/or O₃, acquired from plants growing in monoculture, may not be transferred to plants grown under interspecific competition as typically found in the field.     

Competitive interaction in plant populations exposed to supplementary ultraviolet-B radiation

Summary Changes in plant growth and competitive balance between pairs of competing species were documented as a result of supplementary ultraviolet-B radiation (principally in the 290–315 nm waveband) under field conditions. This component of the terrestrial solar spectrum would be intensified if the atmospheric ozone layer were reduced. A method for calculating and statistically analyzing relative crowding coefficients was developed and used to evaluate the competitive status of the species pairs sown in a modified replacement series. The effect of the supplementary UV-B irradiance was generally detrimental to plant growth, and was reflected in decreased leaf area, biomass, height and density as well as changes in competitive balance for various species. For some species, interspecific competition apparently accentuated the effect of the UV-B radiation, while more intense intraspecific competition may have had the same effect for other species. A few species when grown in a situation of more severe mutual interspecific competition exhibited enhanced growth under the UV-B radiation treatment. This, however, was usually associated with a detrimental effect of the radiation on its competitor and thus was likely the result of its improved competitive circumstance rather than a beneficial physiological effect of the radiation.     

Interactions amongst species in a guild of subtidal benthic herbivores

Summary The subtidal coralline flats of northeastern New Zealand support a characteristic guild of grazing herbivores. The most important members of this guild are an echinometrid echinoid, patellid, turbinid and trochid gastropods. Densities of these herbivores fluctuate through time. Interactions within and among the different species of echinoids and gastropods were investigated experimentally. Different combinations of species were caged at densities up to 5 times that of ambient for a 24 week period in an experiment designed to differentiate between intra- and interspecific competition.The echinoidEvechinus chloroticus and the turbinid gastropodCookia sulcata exhibited reduced mean dry weight with increasing intraspecific densities. There was little evidence of density-related mortality in these species. The limpetCellana stellifera showed comparatively large losses of weight and enhanced mortalities in intraspecific experimental treatments but this was not related to density.Investigation of interspecific interactions amongstEvechinus andCookia revealed no evidence of a negative influence of one species on the other. In terms of dry weight,Cookia was indifferent to the presence ofEvechinus, andEvechinus benefited in the reciprocal interaction.Cookia also enjoyed an enhanced mean dry weight when in the presence ofCellana compared to the equivalent intraspecific treatments. There were no coherent trends in proportional mortality in any treatments with enhanced interspecific densities. Cellana, in the presence ofCookia, exhibited a dramatic decrease in mortality rate and increase in mean dry weight. The presence of the turbinid gastropod was clearly beneficial to the limpet when compared to the intraspecific treatments with enhanced intraspecific densities and the control cages containingCellana at ambient density. We suggest that subtidal areas constitute poor habitats for limpets in the absence of agents such asCookia which may provide or maintain suitable sites for attachment and grazing.For the combinations of densities and species investigated there was a consistent trend towards positive interspecific interactions. It seems unlikely that at the sites investigated interspecific competition could act to restrict distributions, or limit abundances of species.     

Nutrient addition does not increase the benefits of clonal integration in an invasive plant spreading from open patches into plant communities

• One benefit of clonal integration is that resource translocation between connected ramets enhances the growth of the ramets grown under stressful conditions, but whether such resource translocation reduces the performance of the ramets grown under favourable conditions has not produced consistent results. In this study, we tested the hypothesis that resource translocation to recipient ramets may reduce the performance of donor ramets when resources are limiting but not when resources are abundant.
• We grew Mikania micrantha stolon fragments (each consisting of two ramets, either connected or not connected) under spatially heterogeneous competition conditions such that the developmentally younger, distal ramets were grown in competition with a plant community and the developmentally older, proximal ramets were grown without competition. For half of the stolon fragments, slow‐release fertiliser pellets were applied to both the distal and proximal ramets.
• Under both the low and increased soil nutrient conditions, the biomass, leaf number and stolon length of the distal ramets were higher, and those of the proximal ramets were lower when the stolon internode was intact than when it was severed. For the whole clone, the biomass, leaf number and stolon length did not differ between the two connection treatments. Connection did not change the biomass of the plant communities competing with distal ramets of M. micrantha.
• Although clonal integration may promote the invasion of M. micrantha into plant communities, resource translocation to recipient ramets of M. micrantha will induce a cost to the donor ramets, even when resources are relatively abundant.