Transcriptome analysis of intraspecific competition in Arabidopsis thaliana reveals organ-specific signatures related to nutrient acquisition and general stress response pathways.

ABSTRACT: BACKGROUND: Plants are sessile and therefore have to perceive and adjust to changes in their environment. The presence of neighbours leads to a competitive situation where resources and space will be limited. Complex adaptive responses to such situation are poorly understood at the molecular level. RESULTS: Using microarrays, we analysed whole-genome expression changes in Arabidopsis thaliana plants subjected to intraspecific competition. The leaf and root transcriptome was strongly altered by competition. Differentially expressed genes were enriched in genes involved in nutrient deficiency (mainly N, P, K), perception of light quality, and responses to abiotic and biotic stresses. Interestingly, performance of the generalist insect Spodoptera littoralis on densely grown plants was significantly reduced, suggesting that plants under competition display enhanced resistance to herbivory. CONCLUSIONS: This study provides a comprehensive list of genes whose expression is affected by intraspecific competition in Arabidopsis. The outcome is a unique response that involves genes related to light, nutrient deficiency, abiotic stress, and defence responses.

Changes in reproductive investment with altitude in an alpine plant

Aims: In perennial species, the allocation of resources to reproduction results in a reduction of allocation to vegetative growth and, therefore, impacts future reproductive success. As a consequence, variation in this trade-off is among the most important driving forces in the life-history evolution of perennial plants and can lead to locally adapted genotypes. In addition to genetic variation, phenotypic plasticity might also contribute to local adaptation of plants to local conditions by mediating changes in reproductive allocation. Knowledge on the importance of genetic and environmental effects on the trade-off between reproduction and vegetative growth is therefore essential to understand how plants may respond to environmental changes. Methods: We conducted a transplant experiment along an altitudinal gradient from 425 m to 1921 m in the front range of the Western Alps of Switzerland to assess the influence of both altitudinal origin of populations and altitude of growing site on growth, reproductive investment and local adaptation in Poa alpina. Important findings: In our study, the investment in reproduction increased with plant size. Plant growth and the relative importance of reproductive investment decreased in populations originating from higher altitudes compared to populations originating from lower altitudes. The changes in reproductive investment were mainly explained by differences in plant size. In contrast to genetic effects, phenotypic plasticity of all traits measured was low and not related to altitude. As a result, the population from the lowest altitude of origin performed best at all sites. Our results indicate that in P. alpina genetic differences in growth and reproductive investment are related to local conditions affecting growth, i.e. interspecific competition and soil moisture content.

Tree cover at fine and coarse spatial grains interacts with shade tolerance to shape plant species distributions across the Alps

The role of competition for light among plants has long been recognized at local scales, but its potential importance for plant species' distribution at larger spatial scales has largely been ignored. Tree cover acts as a modulator of local abiotic conditions, notably by reducing light availability below the canopy and thus the performance of species that are not adapted to low-light conditions. However, this local effect may propagate to coarser spatial grains. Using 6,935 vegetation plots located across the European Alps, we fit Generalized Linear Models (GLM) for the distribution of 960 herbs and shrubs species to assess the effect of tree cover at both plot and landscape grain sizes (~ 10-m and 1-km, respectively). We ran four models with different combinations of variables (climate, soil and tree cover) for each species at both spatial grains. We used partial regressions to evaluate the independent effects of plot- and landscape-scale tree cover on plant communities. Finally, the effects on species' elevational range limits were assessed by simulating a removal experiment comparing the species' distribution under high and low tree cover. Accounting for tree cover improved model performance, with shade-tolerant species increasing their probability of presence at high tree cover whereas shade-intolerant species showed the opposite pattern. The tree cover effect occurred consistently at both plot and landscape spatial grains, albeit strongest at the former. Importantly, tree cover at the two grain sizes had partially independent effects on plot-scale plant communities, suggesting that the effects may be transmitted to coarser grains through meta-community dynamics. At high tree cover, shade-intolerant species exhibited elevational range contractions, especially at their upper limit, whereas shade-tolerant species showed elevational range expansions at both limits. Our findings suggest that the range shifts for herb and shrub species may be modulated by tree cover dynamics.

Evolutionary Ecology of Mountain Birch in Subarctic Stress Gradients: Interplay of Biotic and Abiotic Factors in Plant-Plant Interactions and Evolutionary Processes

In nature, variation for example in herbivory, wind exposure, moisture and pollution impact often creates variation in physiological stress and plant productivity. This variation is seldom clear-cut, but rather results in clines of decreasing growth and productivity towards the high-stress end. These clines of unidirectionally changing stress are generally known as ‘stress gradients’. Through its effect on plant performance, stress has the capacity to fundamentally alter the ecological relationships between individuals, and through variation in survival and reproduction it also causes evolutionary change, i.e. local adaptations to stress and eventually speciation. In certain conditions local adaptations to environmental stress have been documented in a matter of just a few generations. In plant-plant interactions, intensities of both negative interactions (competition) and positive ones (facilitation) are expected to vary along stress gradients. The stress-gradient hypothesis (SGH) suggests that net facilitation will be strongest in conditions of high biotic and abiotic stress, while a more recent ‘humpback’ model predicts strongest net facilitation at intermediate levels of stress. Plant interactions on stress gradients, however, are affected by a multitude of confounding factors, making studies of facilitation-related theories challenging. Among these factors are plant ontogeny, spatial scale, and local adaptation to stress. The last of these has very rarely been included in facilitation studies, despite the potential co-occurrence of local adaptations and changes in net facilitation in stress gradients. Current theory would predict both competitive effects and facilitative responses to be weakest in populations locally adapted to withstand high abiotic stress. This thesis is based on six experiments, conducted both in greenhouses and in the field in Russia, Norway and Finland, with mountain birch (Betula pubescens subsp. czerepanovii) as the model species. The aims were to study potential local adaptations in multiple stress gradients (both natural and anthropogenic), changes in plant-plant interactions under conditions of varying stress (as predicted by SGH), potential mechanisms behind intraspecific facilitation, and factors confounding plant-plant facilitation, such as spatiotemporal, ontogenetic, and genetic differences. I found rapid evolutionary adaptations (occurring within a time-span of 60 to 70 years) towards heavy-metal resistance around two copper-nickel smelters, a phenomenon that has resulted in a trade-off of decreased performance in pristine conditions. Heavy-metal-adapted individuals had lowered nickel uptake, indicating a possible mechanism behind the detected resistance. Seedlings adapted to heavy-metal toxicity were not co-resistant to others forms of abiotic stress, but showed co-resistance to biotic stress by being consumed to a lesser extent by insect herbivores. Conversely, populations from conditions of high natural stress (wind, drought etc.) showed no local adaptations, despite much longer evolutionary time scales. Due to decreasing emissions, I was unable to test SGH in the pollution gradients. In natural stress gradients, however, plant performance was in accordance with SGH, with the strongest host-seedling facilitation found at the high-stress sites in two different stress gradients. Factors confounding this pattern included (1) plant size / ontogenetic status, with seedling-seedling interactions being competition dominated and host-seedling interactions potentially switching towards competition with seedling growth, and (2) spatial distance, with competition dominating at very short planting distances, and facilitation being strongest at a distance of circa ¼ benefactor height. I found no evidence for changes in facilitation with respect to the evolutionary histories of plant populations. Despite the support for SGH, it may be that the ‘humpback’ model is more relevant when the main stressor is resource-related, while what I studied were the effects of ‘non-resource’ stressors (i.e. heavy-metal pollution and wind). The results have potential practical applications: the utilisation of locally adapted seedlings and plant facilitation may increase the success of future restoration efforts in industrial barrens as well as in other wind-exposed sites. The findings also have implications with regard to the effects of global change in subarctic environments: the documented potential by mountain birch for rapid evolutionary change, together with the general lack of evolutionary ‘dead ends’, due to not (over)specialising to current natural conditions, increase the chances of this crucial forest-forming tree persisting even under the anticipated climate change.

Effect of plant row spacing and herbicide use on weed aboveground biomass and corn grain yield

The use of narrow plant spacing in corn (Zea mays) has been suggested as a technological alternative to obtain grain yield increases, due to a better use of resources. The regular pattern could diminish intraspecific competition while favoring interspecific competition with weeds. The objective of this study was to analyze the effect of corn row spacing on weed aboveground biomass and corn grain yield. Field experiments were conducted during 2002/2003 and 2003/2004 growing seasons. Three corn hybrids with two-row width (0.70 and 0.35 m) were tested. A greater photosynthetic photon flux density (PPFD) interception with a lower weed aboveground dry matter in narrow row arrangement was obtained. Corn grain yield was greater in the narrow row arrangement than in the wide row spacing. This increase in grain yield was related to a better resource use that allows for a reduced interspecific competition. The use of reduced spatial arrangement appeared to be an interesting alternative to increase both the grain yield potential and corn suppressive ability against weeds in corn dryland production systems.

Effects of weed species competition on the growth of young coffee plants

The effects of competition of seven weed species on the growth of coffee plants were evaluated under greenhouse conditions. Thirty days after coffee seedling transplantation into 12 L pots with soil level area of 6.5 dm², weeds were transplanted into or sown in those pots, at densities of 0, 1, 2, 3, 4 and 5 plants per pot. Competition or weedy periods from weed transplantation or emergence to plant harvesting, at weed pre-flowering stage, were: 77 days - Bidens pilosa, 98 days - Brachiaria decumbens, 180 days - Commelina diffusa, 82 days - Leonurus sibiricus, 68 days - Nicandra physaloides, 148 days - Richardia brasiliensis and 133 days - Sida rhombifolia. Coffee plant height, stem diameter, leaf number and shoot dry matter were determined. Effects of competition by N. physaloides and S. rhombifolia against coffee plants were among the lowest, since only a slight decrease in all the characteristics evaluated in coffee plants was observed. The other weed species caused severe decrease in growth, mainly with increasing weed plant densities. Competition degree was found to depend on weed species and density.

Growth and nutrient concentration in coffee root system under weed species competition

The effects of competition of six weed species on growth, nutrient concentration and nutrient content of coffee plant root system under greenhouse conditions were evaluated. Thirty days after coffee seedling transplantation into 12 L pots with soil level area of 6.5 dm². Weeds were transplanted or sowed in these pots, at densities of 0, 1, 2, 3, 4 and 5 plants per pot. The duration of competition (or weedy periods) from weed transplantation or emergence until plant harvesting, at the weed preflowering stage, were (in days): 77 (Bidens pilosa), 180 (Commelina diffusa), 82 (Leonurus sibiricus), 68 (Nicandra physaloides), 148 (Richardia brasiliensis) and 133 (Sida rhombifolia). Dry matter of coffee plants was linearly reduced with increasing B. pilosa and S. rhombifolia density, with pronounced effect of B. pilosa. C. diffusa was the only weed species whose increasing density in the pots did not diminish crop root dry matter. L. sibiricus, N. physaloides and R. brasiliensis reduced root dry matter of coffee plants by 75, 52 and 47%, respectively, as compared to the weed-free treatment, regardless of weed density. Under competition, even though weed species showed lower macronutrient concentration in the roots (except for P), they accumulated 4.2 (N), 12.3 (P), 4.3 (K), 5.5 (Ca), 7.6 (Mg) and 4.4 (S) times more nutrients in the roots than the coffee plants. Crop and weed nutrient concentration, as well as competition degrees greatly varied depending on both weed species and densities.

Growth and yield of corn grain and green ear in competition with weeds

Studies on plant growth are interesting because they provide explanations for the factors that influence yield in various crops. The objective of this work was to evaluate growth and yield in corn cultivar AG1051, when in competition with weeds. Cultivar AG 1051 was submitted to two groups of treatments: weed control, and sampling periods for dry biomass evaluation. The weed control treatments consisted of hoeing (two hoeings performed at 20 and 40 days after sowing) and no hoeing. Sampling periods consisted of collecting the above-ground part and roots of corn every fifteen days, until 105 days after sowing (DAS); the first sampling was performed 30 DAS. A completely randomized block design with ten replicates was used. For the characteristics evaluated in a single season, statistical analyses were carried out as a random block experiment. For the characteristics evaluated in several periods, statistical analyses were carried out as random blocks with split-plots (weed control assigned to plots). Fourteen weed species, unevenly distributed throughout the experimental area, were the most important. The growth observed for the above-ground part and root system of corn was 30% smaller in the non-hoed plots, compared to the hoed plots. Lack of weed control increased dry matter of the above-ground part of the weeds and reduced the number of unhusked and husked marketable green ears by 23% and 49%, respectively. Grain yield reduction caused by lack of weed control reached 38%.

Green ear yield and grain yield of maize cultivars in competition with weeds

The reduction in herbicide use is one of the greatest interests for modern agriculture and several alternatives are being investigated with this objective, including the adoption of cultivars that suppress weeds. The objective of this study was to verify if maize cultivars develop differently, in competition with weeds, to produce green ears and grain. Randomized complete block design was used, with split-plots and five replications. Cultivars DKB 390, DKB 466, DKB 350, AG 7000, AG 7575 and Master, were evaluated in the plots, without weeding and two weedings (at 22 and 41 days after sowing) in sub plots. Twenty-one species were identified in the experimental area, the most frequent being Gramineae (Poaceae), Euphorbiaceae, Leguminosae (Fabaceae) and Convolvulaceae species. There was no difference in the dry biomass above-ground part of the weeds in the plots of the evaluated cultivars. The cultivars behaved similarly in treatments with or without hoeing, except for plant height and ear height evaluations. Without hoeing, plant height increased in cultivar DKB 390, while plant height and ear height decreased in cultivar AG 7575. In the other cultivars, these traits did not change under weed control. The presence of weeds decreased the values of all traits employed to assess green corn yield, with the exception of the total number of green ears and grain yield.

Corn growth and yield in competition with weeds

Although labor is intensive, evaluating the growth of crops may allow a better understanding of crop performance, including the reasons why certain cultivars can compete better with weeds. This study aims at evaluating growth, green ear yield, and grain yield in corn when in competition with weeds. Cultivars AG 1051 and BRS 106 were grown with (two hoeings, at 20 and 40 days after sowing) or without weed control. In order to evaluate crop growth, six collections of the above-ground part and the root system of corn were performed, every 15 days, with the first collection made 30 days after sowing. A randomized complete block design was adopted, with split-split plots (weed control in plots, cultivars in subplots, and collections in sub-subplots) and ten replicates. Eighteen weed species were found in the experiment area. Increased values of corn leaf area, above-ground part and root system, due to plant age function, were smaller in non-hoed plots than in hoed plots and were dependent upon cultivar. The lack of weed control increased dry matter of weeds aboveground part and decreased green ear yield and grain yield. Cultivar AG 1051 had higher increases in leaf area, above-ground part of the plant and root system, due to plant age function, and controlled weeds better than cultivar BRS 106. In addition, cultivar AG 1051 was superior to other cultivars with respect to most traits used for green corn yield and grain yield assessment.

Competition between drought-tolerant upland rice cultivars and weeds under water stress condition

The objective of this study was to evaluate the competitiveness of two cultivars of upland rice drought-tolerant, cultured in coexistence with weed S. verticillata, under conditions of absence and presence of water stress. The experiment was conducted in a greenhouse at the Experimental Station of the Universidade Federal de Tocantins, Gurupi-TO Campus. The experimental design was completely randomized in a factorial 2 x 2 x 4 with four replications. The treatments consisted of two rice cultivars under two water conditions and four densities. At 57 days after emergence, were evaluated in rice cultivars and weed S. verticillata leaf area, dry weight of roots and shoots and total concentration and depth of roots. Was also evaluated in rice cultivars, plant height and number of tillers. Water stress caused a reduction in leaf area, the concentration of roots and vegetative components of dry matter (APDM, and MSR MST) of rice cultivars and Jatoba Catetão and weed S. verticillata. The competition established by the presence of the weed provided reduction of all vegetative components (MSPA, and MSR MST) of cultivars and Jatoba Catetão. It also decreased the number of tillers, the concentration of roots and leaf area. At the highest level of weed competition with rice cultivars, a greater decrease in vegetative components and leaf area of culture, regardless of water conditions.

Effect of plant density and seed position on mother plant on physiological characteristic of cocklebur (Xanthium strumarium) seeds

Experiments were conducted in 2010 to determine the influence of plant density and seed position on the mother plant on seed physiological characteristics of cocklebur (Xanthium strumarium). Cocklebur burs were collected in fall of 2010 from Research Farm of University of Agricultural Sciences and Natural Resources of Gorgan, Iran. The experiment was established as factorial arrangement using a completely randomized design with three replications. The factors included different densities of cocklebur (0, 2, 4, 6 and 8 plant m-2) and the top and bottom parts of the canopy. Non dormant seeds were used for determining cardinal temperatures and tolerance to salinity and drought stresses. Base, optimum and ceiling germination temperatures were estimated between 7.09 to 12.33, 32 to 35 and 44 to 45 respectively in different treatments. Salinity stress up to 300 Mm and osmotic potential 8 bar inhibited the germination completely. Comparison of base temperatures and sigmoid equation coefficients showed that seeds produced in the top had higher germination than those that produced at the bottom of the mother plant. It seems plant densities through seed position on the mother plant affect seed quality. Likewise changes of light quality and quantity in shade environment increased seed dormancy in matured seeds. Shade environment affect seed germination on mother plant that increased dormancy of seeds maturing under shade be an adaptive response that reduces the probability of germination of offspring under unfavorable (shade, competitive) conditions.

Wild onion (Asphodelus tenuifolius) competition in rainfed chickpea-chickpea cropping system

Chickpea yield potential is limited by weed competition in typical chickpea growing areas of Pakistan where zero tillage crop grown on moisture conserved from rains received during the months of September and August. The objective of this work was to evaluate the growth and yield characteristics of chickpea grown in coexistence with increasing densities of wild onion (Asphodelus tenuifolius). The experiment was comprised of six density levels viz. zero, 20, 40, 80, 160 and 320 plants m-2 of A. tenuifolius. A decrease in chickpea primary and secondary branches per plant, pods per plant, seeds per pod, 100-seed weight and seed yield was observed due to more accumulation of dry matter per increasing densities of A. tenuifolius. The increase in A. tenuifolius density accelerated chickpea yield losses and reached the maximum values of 28, 35, 42, 50, 58 and 96% at 20, 40, 80, 160 and 320 A. tenuifolius plants m-2, respectively. The yield loss estimation model showed that chickpea losses with infinite A. tenuifolius density were 60%. Yield reduction could be predicted by 2.52% with increase of one A. tenuifolius plant m-2. It is concluded that A. tenuifolius has a strong influence on chickpea seed yield and showed a linear response at the range of densities studied.

Grass Weeds Interfering with Eucalypt: Effects of the Distance of Coexistence on the Initial Plant Growth

Two experiments were carried out to evaluate the initial plant growth of Eucalyptus urograndis growing in coexistence with Urochloa decumbens and U. ruziziensis. In 100-L box, one plant of U. decumbens or U. ruziziensis grew in coexistence with one plant of E. urograndis clones C219H or H15, respectively, in the distances of 0, 5, 10, 15, 20, 25, 30, 35, and 40 cm from the crop. After 30, 60, 90 (both clones), and 150 days (just for H15), growth characteristics were evaluated. Plants of both clones, growing in weed-free situations, showed a better growth and development than plants that grew in weedy situations, independently of the distance, having the highest plant height, stem diameter, dry mass of stem, and dry mass of leaves. As the same way, the number of branches, number of leaves, and leaf area of the clone C219H were similarly affected. Urochloa ruziziensis reduced the dry mass accumulation of stem and leaves by the rate of 0.06 and 0.32 g per plant, respectively, per each centimeter growing nearest to the crop, while U. decumbens reduced by 0.03 and 0.14 g per plant. The interference of U. decumbens and U. ruziziensis with E. urograndis is more intense when weedy plants grow in short distances from the crop.

Wheat Yield Loss in a Two Species Competition with Emex australis and Emex spinosa

Emex australis and E. spinosa are significant weed species in wheat and other crops. Information on the extent of competition of the Emex species will be helpful to access yield losses in wheat. Field experiments were conducted to quantify the interference of tested weed densities each as single or mixture of both at 1:1 on their growth and yield, wheat yield components and wheat grain yield losses in two consecutive years. Dry weight of both weed species increased from 3-6 g m-2 with every additional plant of weed, whereas seed number and weight per plant decreased with increasing density of either weed. Both weed species caused considerable decrease in yield components like spike bearing tillers, number of grains per spike, 1000-grain weight of wheat with increasing density population of the weeds. Based on non-linear hyperbolic regression model equation, maximum yield loss at asymptotic weed density was estimated to be 44 and 62% with E. australis, 56 and 70% with E. spinosa and 63 and 72% with mixture of both species at 1:1 during both year of study, respectively. It was concluded that E. spinosa has more competition effects on wheat crop as compared to E. australis.