Competition between genotypes within a plant population can result in the displacement of the least competitive by more competitive genotypes. Although evolutionary processes in plants may occur over thousands and millions of years, it has been suggested that changes in key fitness traits could occur in as little as decades, with herbicide resistance being a common example. However, the rapid evolution of complex traits has not been proven in weeds. We hypothesized that changes in weed growth and competitive ability can occur in just a few years because of selection in agroecosystems. Seed of multiple generations of a single natural population of the grassy weed giant foxtail (Setaria faberi Herrm.) were collected during 34 yr (i.e., 1983 to 2017). Using a “resurrection” approach, we characterized life-history traits of the different year-lines under noncompetitive and competitive conditions. Replacement-series experiments comparing the growth of the oldest year-line (1983) versus newer year-lines (1991, 1996, 1998, 2009, and 2017) showed that plant competitive ability decreased and then increased progressively in accordance with oscillating selection. The adaptations in competitive ability were reflected in dynamic changes in leaf area and biomass when plants were in competition. The onset of increased competitive ability coincided with the introduction of herbicide-resistant crops in the landscape in 1996. We also conducted a genome-wide association study and identified four loci that were associated with increased competitive ability over time, confirming that this trait changed in response to directional selection. Putative transcription factors and cell wall–associated enzymes were linked to those loci. This is the first study providing direct in situ evidence of rapid directional evolution of competitive ability in a plant species. The results suggest that agricultural systems can exert enough pressure to cause evolutionary adaptations of complex life-history traits, potentially increasing weediness and invasiveness.

The design and analysis of competition experiments should be based on an unambiguous objective. Recent criticisms of particular designs have been made without reference to objectives and may be misleading. Three common designs are discussed: additive, replacement series, and response surface. Additive designs are well suited to agronomic objectives; replacement series are useful for comparing pairs of species; response surface designs can be used for most objectives but may be unnecessarily complex. The published criticisms of additive and replacement series designs are argued to be acceptable limitations within the bounds of the objectives for which they are used. Concerns about these designs confounding density and proportion are irrelevant to the objectives for which they are most suited. The continued use of multiple comparison tests is argued to be illogical. Regression approaches to analysis are more relevant, many non-linear equations are now easy to fit to data and these can be used without the need for linearization. However, there are various pitfalls not adequately reported to date. In particular, error structures need to be checked carefully and over-elaborate equations should be avoided.

Greenhouse experiments in central Texas assessed the relative importance of above- and belowground interactions of semidwarf Mit wheat and Marshall ryegrass during vegetative growth. One experiment used partitions to compare the effect of no (controls), aboveground only, belowground only, and full interaction for 75 d after planting (DAP) one wheat and nine ryegrass plants in soil volumes of 90, 950, and 3,800 ml. The results with the different soil volumes were similar. Wheat growth in the aboveground interaction only did not differ from controls. However, the full or belowground only interaction of wheat with ryegrass reduced wheat height, leaf number, tillering, leaf area, percent total nonstructural carbohydrates in shoot, and dry weights of leaves, stems, and roots 45 and 75 DAP compared to controls. Wheat in full and belowground interaction only did not differ from one another in growth. A replacement series experiment of 56 d also showed that the competitive advantage of ryegrass was relatively greater in root than in shoot growth. No allelopathic response of wheat to ryegrass occurred. While the tallness of the semidwarf wheat minimized aboveground interference by ryegrass, the root growth of the thinner and more fibrous roots of ryegrass greatly enhanced its belowground competitiveness.

Growth characteristics and competitive relationships of cotton and three perennial weeds (johnsongrass, purple nutsedge, and yellow nutsedge) were investigated at the USDA Cotton Research Station in Shafter, CA. In growth analysis experiments, yellow nutsedge and johnsongrass had highest values for height, biomass, leaf area production, growth rate, and photosynthetic efficiency. The weeds exhibited greater overall resource use and production efficiency than cotton, while cotton attained greater leafiness and canopy closure than weeds over the 10-wk experiment. Linear correlation indicated that most growth variables were significantly correlated with aggressivity (competitiveness); however, all correlation coefficients were less than 0.80. Stepwise multiple regression, using aggressivity as the dependent variable, defined 4 growth variables out of 12 that best described competitiveness. These were unit leaf rate (ULR), height, relative growth rate (RGR), and initial propagule weight (PWT). Thus, parameters of light utilization (ULR, height) and early establishment (RGR, PWT) were best predictors of competitive success in this system.

In replacement experiments in the greenhouse, plant relative yield (PRY) of both eastern black nightshade and tomato increased as the proportion of nightshade plants increased in the pots, indicating that nightshade is less competitive than tomato. In field studies tomato yield was reduced by two-thirds if three nightshade plants m–1 of row were allowed to grow with tomato more than 6 weeks following tomato establishment The percent marketable fruit decreased linearly from 73% with no nightshade to 49% when nightshade were present for 12 weeks. When nightshade and tomato were transplanted together, tomato yield was 9000 kg ha–1 and 49% of the fruit was marketable, while tomato yields were 30 000 kg ha–1 and 70% of the fruit was marketable when nightshade was established 9 weeks after tomato planting.

Wild, ball, and dog mustard growth and development were investigated by mathematical growth analysis in a greenhouse experiment. Plant height and total plant biomass over the growth period followed the trend wild mustard > ball mustard > dog mustard. Dog mustard plants had lower leaf areas than either wild or ball mustard. In a replacement series experiment, wild mustard was more competitive than either ball or dog mustard, and ball mustard was more competitive than dog mustard.

Greenhouse, growth chamber, and laboratory studies were conducted to determine anatomical and morphological characteristics and cultural practices limiting the distribution of cogongrass, torpedograss, and johnsongrass in the United States. Cogongrass did not produce axillary buds along most of the rhizome nor regenerate when apical six-node-long rhizome segments were buried deeper than 8 cm. Both torpedograss and johnsongrass produced axillary buds along the entire lengths of their rhizomes. Torpedograss shoot emergence decreased at burial depths between 8 and 16 cm. Shoot emergence from johnsongrass rhizomes was not affected by burial as deep as 16 cm. Rhizomes of all three species were tolerant of desiccation. Cogongrass grew better in soil at pH 4.7 than in soil at pH 6.7, whereas torpedograss and johnsongrass grew equally well in either pH. It is postulated that cogongrass spread is limited by lack of axillary bud formation on most of the rhizome and the inability of rhizomes to send up new shoots if buried deeper than 8 cm. These factors could account for the intolerance of cogongrass to cultivation. Torpedograss appears to spread only vegetatively due to the lack of viable seed production.

The relative competitiveness of triazine-resistant (TR) and -susceptible (TS) common waterhemp, as a component of fitness, was assessed using TR and TS biotypes collected from the same field in Fillmore County, Nebraska. Competition between biotypes was evaluated under greenhouse conditions using a set of replacement series experiments with five ratios of TR:TS (1:0, 3:1, 1:1, 1:3, and 0:4) and four plant densities (50, 100, 150, and 300 plants m−2). No difference in final individual plant weight between TR and TS plants was observed in the absence of interbiotypic competition at all plant densities examined. Interbiotypic competition was indicated by ANOVA of plant weight and modified relative crowding coefficients (RCC) for plant weight, height, and volume. Replacement series diagrams showed TS plants were more competitive than TR plants, as measured by plant weight at densities of 50, 100, and 150 plants m−2. TS and TR plants were equally competitive at 300 plants m−2. Individual TS plant weight for 50, 100, and 150 plants m−2 was greatest at the 3:1 ratio (TR:TS) and decreased as the proportion of susceptible plants increased. Individual TR plant weight for 50, 100, and 150 plants m−2 was greatest at the 1:0 ratio and decreased as the TR proportion decreased. Lack of interbiotypic competition at 300 plants m−2 and decreasing TS plant weight with increasing ratios suggest intrabiotypic competition. RCC values for volume and height were calculated weekly during the experiment and indicated the competitive advantage of TS plants was constant from transplanting through harvest.

Destructive growth analysis of field replacement series experiments with mayweed chamomile and dry field peas was used to determine the competitive relationship between the two species. Mayweed chamomile produced similar amounts of leaf area and dry weight per plant in a dry year and a wet year. On the other hand, dry field peas produced 20% more leaf area and 100% more dry weight per plant in the wet year compared to the dry year. Height, leaf area, and dry weight of peas reached maximum between bloom and pod-set, and then declined. Height and dry weight of mayweed chamomile increased steadily throughout the growing season. Mayweed chamomile leaf area reached a maximum at the beginning of flower stem elongation. Initially, the relative growth rate of mayweed chamomile was about three times greater than the rate for peas, but by 40 to 48 d after planting, rates were similar for both species. Relative yields and relative crowding coefficients for dry weight showed that peas were 3 to 20 times more aggressive than mayweed chamomile. Results of these experiments show that mayweed chamomile is a weak competitor against peas, but because it continues to grow after peas senesce, it could interfere with crop harvest.

Three approaches to data analysis were compared to describe competitive interactions between wheat and Italian ryegrass. Replacement series were performed using the two species at total densities of 100, 200, and 400 plants/ m2, and separate monoculture experiments for each species at densities from 33 to 800 plants/m2. Approaches to data analysis included: 1) conventional analysis of replacement series experiments, 2) development of synthetic no-interaction responses from monoculture experiments for comparison with results from mixtures, and 3) responses of the reciprocal yield of individual plants to variation in densities of the two species. Wheat was the superior competitor to ryegrass; however, the three approaches varied in ability to quantify this competitive relationship. The conventional replacement series analysis was least sensitive in describing the influences of either density or proportion on the plant association. The synthetic no-interaction approach provided the most detailed analysis of the influence of proportion on the species interaction. The reciprocal yield approach provided the simplest and most sensitive analysis of the joint influences of density and proportion. The latter approach also provided the most quantitative analysis of the influence of density on the species interaction. Plant density and species proportion are important variables for interpreting the process of plant competition.

Replacement series studies were conducted under controlled conditions to determine the effect of phosphorus (P) rates and population densities on the competitiveness of smooth pigweed and common purslane with lettuce. Densities were 2, 4, and 8 plants per 113 cm2, whereas P rates were 0, 0.4, and 0.8 g PL−1 soil. A P-deficient Histosol (0.3 mg water-extractable P L−1 soil) was used. High P fertility enhanced the competitive ability of lettuce in smooth pigweed-lettuce mixtures. Smooth pigweed was not responsive to P rates. However, luxurious P consumption by smooth pigweed occurred, reducing the amount of the nutrient available for lettuce absorption. In common purslane-lettuce mixtures, the weed was responsive to P rates, increasing its competitive ability, whereas no increase in lettuce competitive ability was observed. Both weed species were more competitive than lettuce. Competition for P appears to be the main mechanism of common purslane interference on lettuce grown in low-P organic soils. Alternative fertilization strategies (i.e., banded applications) may reduce the effect of smooth pigweed on lettuce.

Replacement series experiments were conducted under greenhouse conditions to evaluate effects of population densities and proportions on the intraspecific and interspecific interference of either purple nutsedge or yellow nutsedge with tomato cv. Sunny, under nonlimiting conditions of water and nutrients. When grown with either nutsedge species for 40 d, tomato dry weight per plant increased and dry weight per plant of nutsedge decreased as their relative proportions decreased in mixture. Relative yield analysis indicated tomato is a stronger competitor than either nutsedge species. Both nutsedges appeared to be weak interspecific competitors but strong intraspecific competitors. Attenuated light on a purple nutsedge canopy showed that tomato additions can reduce light received by the weed compared to pure nutsedge stands.

Substitutive (replacement) and partial additive experimental designs, with their underlying models, remain the two most popular techniques in weed-crop competition studies, despite considerable criticism of these approaches in the recent literature. We review standard designs for two-species competition experiments and demonstrate the advantages of a reciprocal yield model applied to data from an additive series experiment, using mixtures of Japanese millet and tomato. A traditional replacement series analysis failed to provide a general model of competition among these two species over several total plant densities, while an application of a reciprocal yield (inverse linear) model to the same data was successful. This technique allows evaluation of the influences of both weed on crop and crop on weed, as well as the partitioning of net competition effects into intra- and interspecific components. One Japanese millet plant was competitively equivalent to 3.7 tomato plants, as measured by effects on tomato biomass, while one tomato plant was equivalent to 0.14 Japanese millet plants, as measured by effects on millet biomass. Skewness of per plant biomass distribution is shown to be a result but not an unambiguous measure of competition. Expansion of a reciprocal yield model to mixtures of more than two species is illustrated using three species of duckweed. While some caution is recommended, the reciprocal yield analysis applied to data from appropriately designed experiments is a substantial improvement over more traditional methods.

Intraspecific and interspecific interference effects on growth, gas exchange, and water potential of wild oat and spring barley were measured under greenhouse conditions using a 1:1.06 barley to wild oat replacement series. Intraspecific barley interference affected barley growth more than interspecific wild oat interference. Interspecific wild oat interference with barley reduced wild oat growth more than intraspecific interference. Wild oat plant height surpassed barley plant height near barley anthesis. Growth and gas exchange of barley and wild oat responded the same to short-term water stress.

A greenhouse experiment compared the vegetative growth in pure cultures and mixtures of winter Triticum aestivum cultivar ‘Mit’ and Lolium multiflorum cultivar ‘Marshall’ in continuously watered controls and drought treatments. Control L. multiflorum in pure culture 14 wk after planting produced more leaf area, tillers, and dry weights of stem and root than control T. aestivum in pure culture. The greater seed size, larger initial leaf area, and height allowed T. aestivum to produce greater final leaf area and dry stem weight in control mixtures than L. multiflorum. Watering following drought shifted the relative performance of the two species in pure cultures and mixtures compared to controls. The ability of T. aestivum to maintain a greater leaf expansion rate during drought and a greater leaf area afterward than L, multiflorum allowed T. aestivum to attain greater growth than L. multiflorum in pure cultures exposed to temporary drought followed by watering. Conversely, drought and its relief enhanced the relative competitiveness of L. multiflorum compared to controls in mixtures with T. aestivum. During 4 wk of watering following the drought, L. multiflorum in mixtures grew vigorously and was similar to T. aestivum in all measures except in height and dry stem weight. Thus, L. multiflorum was similar in root growth with T. aestivum in control and drought mixtures and had its aboveground competitiveness amplified by the cycle of drought and watering in this study. There was no evidence of an allelopathic interaction between the two species.

Experiments were conducted to determine the productivity and the competitive ability of a large crabgrass accession (PW2) resistant to acetyl-coenzyme A carboxylase (ACCase) inhibitors relative to a susceptible accession (A310). Under noncompetitive conditions in the field, shoot dry biomass and plant height over time were similar for the two accessions. Leaf area was slightly greater for A310 than for PW2. The instantaneous relative growth rate was similar for the accessions. The instantaneous net assimilation rate was slightly greater for A310 than for PW2; however, the instantaneous leaf area ratio was slightly greater for PW2 than for A310. At maturity, the seed yield was similar for the accessions. The inter-accession competitive ability of PW2 and A310 plants in the field was similar, based on shoot dry biomass or seed yield. In the greenhouse, interspecific competitive ability of the accessions with corn was similar, based on shoot dry biomass or seed yield. These results suggest that resistance to ACCase inhibitors is not associated with reduced fitness of the PW2 accession.

Early weed and crop interference was examined in a greenhouse pot experiment combining aspects of additive, substitutive, and diallel designs. Cotton (Gossypium hirsutum L. 'Stoneville 213′) and four weeds {purple nutsedge (Cyperus rotundus L.), large crabgrass [Digitaria sanguinalis (L.) Scop.], velvetleaf (Abutilon theophrasti Medic.) and prickly sida (Sida spinosa L.)} were grown singly and in all possible combinations of pairs including reciprocals for 5 weeks in 10-cm plastic pots. After 5 weeks large crabgrass was the tallest species in all combinations except when grown with purple nutsedge. Purple nutsedge, however, had more leaf area and greater dry weight than all other species. Height, leaf area, and dry weight were lowest in prickly sida. Mean relative yields in mixtures (a measure of competitive ability) were 1.90, 1.15, 1.13, 0.86, and 0.41 for purple nutsedge, large crabgrass, cotton, velvetleaf, and prickly sida, respectively, when grown with the other four species as associates. Conversely, purple nutsedge and cotton as associates depressed mean relative yields of competing species to 0.59 and 0.94. Large crabgrass, velvetleaf, and prickly sida as associates increased mean relative yields of competing species to 1.17, 1.18, and 1.55, respectively. Analysis of these data and replacement-series diagrams indicated that purple nutsedge was the most competitive, prickly sida the least, and the other species were intermediate and approximately equivalent in competitive ability.

Replacement series and growth analysis experiments under greenhouse and field conditions quantified and explained the competitive ability of wild oats and barley. Barley was a stronger competitor than wild oats under greenhouse and field conditions. The reciprocal yield approach showed that barley's intraspecific competition was 7.3 times greater than its interspecific competition with wild oats when calculated on a dry weight per plant basis. When leaf area per plant was the yield variable, barley's intraspecific competition was only 2.4 times greater than its interspecific competition. The difference was explained by wild oats' higher leaf area ratio. Barley had a greater leaf area, root and shoot biomass, absolute growth rate, and shoot-root ratio than wild oats, but wild oats' leaf area ratio was always higher. No differences were detected in relative growth rate and net assimilation rate.

In a replacement series study, barley was more competitive than green foxtail and wild oat at 28/22 and 22/16 C. Wild oat was more competitive with green foxtail at 22/16 C than at 28/22 C. Maximum green foxtail dry weight and leaf area was produced at 28/22 C. As temperature increased, maximum dry weight and leaf area occurred earlier during plant growth. Similarly, maximum wild oat leaf area occurred earlier with increasing temperature, but the greatest leaf area was observed at 16/10 C. Leaf area ratio (LAR) and relative growth rate (RGR) of wild oat did not differ markedly between temperature regimes. Green foxtail LAR was higher while RGR was lower at 16/10 C than at 22/16 or 28/22 C. Under the 16/10 C regime green foxtail produced little mature seed. Greatest seed numbers were produced at 28/22 C. Wild oat produced mature seed under all temperature regimes with the greatest seed numbers produced at 22/16 C.

The relative competitive abilities of yellow starthistle accessions that are resistant (R) and susceptible (S) to picloram were compared using a replacement series experiment. With no herbicide treatment, total shoot dry weights at vegetative and early reproductive stages of plant growth were similar for the two accessions, although S plants accumulated more total shoot dry weight by the late reproductive stage, mainly as a result of a greater contribution of vegetative growth. Without herbicide, relative yield of total biomass or reproductive structures did not differ from theoretical competitive equivalence at any accession ratio, thereby indicating that interaccession interference was similar. For picloram-treated plants, R plants accumulated more total, vegetative, and reproductive dry weight than did S plants at the early and late reproductive stages, and there was no difference between S and R plants at the vegetative growth stage. Seed production by R plants was 10-fold greater than that observed in S plants, but seed size remained unchanged, regardless of accession ratio. With herbicide present, the relative yield of S plants differed from theoretical competitive equivalence as S:R accession ratios decreased, but relative yield of R plants did not. Therefore, only in the presence of picloram will R plants have a competitive advantage over S plants. Some of the progeny from mixed populations of S and R plants that were cross-pollinated, even at low R frequency (25%), expressed resistance to picloram.